Background

Melanin is the most common pigment in plumage. Two classes of melanin form the basis of the colours typically seen in birds namely eumelanins which is responsible for black and brown hues and phaeomelanins which is responsible for reddish-brown colours, but birds often possess a mixture of the two melanin types. These colours are not strongly infuenced by environmental conditions but instead these pigments are produced in cells (Paxton 2009). It is predicted that the downy patterns of chicks have adapted to match habitat. The typical combination of lighter and darker chick plumage on chicks is a matter of pigmentation and dilution of pigment (lighter coloured plumage) respectively.

Plumage colouration and patterning can be used to infer evolutionary relationships (Vázquez and Gittleman 1998; Paxton 2009), and chick plumage patterns and colours have often been used to test phylogenetic relationships among bird taxa (Whetherbee 1957; Bertelli et al.2002). Chicks of related species generally appear particularly similar, which is ascribed to the lack of known selective pressures for rapid divergence in chick plumage patterns during speciation, and therefore lends itself to a phylogenetic analysis that reveals evolutionary relationships. Chick plumage patterns can therefore often provide more valuable clues to phylogenetic relationships than adult plumage patterns (Milstein and Wolff 1987;Johnsgard 2008).

Gamebird chicks are highly cryptic, since they are fightless and avoid predation on the ground (Milstein and Wolff 1987). It would therefore make sense that inheritable traits and crypsis would interact, since natural selection also plays a role (Peichel 2014). Habitat matching, which refers to the consequence of natural selection where plumage colour and pattern matches the colour and texture of occupied habitats, enable birds to avoid detection and capture by predators (Short 1967;Frost 1975; Merilaita and Lind 2005; Stevens and Merilaita 2008).

Chick plumage has been found to have phylogenetic signal to delineate families and to differentiate between sub-families and species. Chick plumage studies indicate that the cassowaries (Casuaridae) and emu (Dromiceiidae) are closely related, and suggest that ostriches(Struthioniformes), cassowaries and emu (Casuariiformes) may be closely related (Jehl 1971). Bertelli et al.(2002) reviewed the chick plumage studies by Jehl (1971)and disputed the division of the tinamous (Tinamidae)into sub-families. In their response Bertelli et al. (2002)acknowledged that forest-dwelling chicks are generally dark in colour with a distinctive forehead, while the steppe tinamou species are striped. While this was used by Jehl (1971) as a cue to describe two sub-families, Bertelli et al. (2002) argued that these chicks possess plesiomorphic character states found in the family. In other words, the tinamous remain a monophyletic family.

外交政策传统反映一国长期、稳定的对外行为的偏好与倾向性。缅甸人在确立中立外交政策的过程中，所体现出的不安全感、地缘心态、对大国的防范、对自身的定位与认识，是构成中立外交传统的基本要素，已内化为缅甸传统的政治文化，也是外界观察其外交政策趋势的基本维度。

Phylogenetic analysis of 157 characters of definitive plumages and soft parts, chick plumage, tracheae, and non-tracheal skeletons of 59 Anatini provided a phylogenetic hypothesis of high consistency (Livezey 1991).The heads of duckling species show different patterns which are particularly evident from the side of the face(laterally). Chicks of closely related species, such as Anas fulvigula, A. castanea and A. acuta are distinguishable based on head patterns (Livezey 1991). Duck chicks are the only group of non-passerine birds studied to date that show differentiation at the species level based on chick plumage. This is unlike the results arrived at by Jehl(1971) and Livezey (1991), whose chick plumage studies reached only sub-family level.

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The chick plumage colour patterns of grouse (Tetraoninae) vary within the range of patterns revealed by other Phasianids, found for example in Perdix perdix and Colinus virginianus. Indeed, a significant pattern identified within the sub-family Tetraoninae included a variegated back, with longitudinal rufous stripes stretching forward onto the crown, a variety of facial marks and distinctive yellowish or greyish white under parts. The Tetraoninae possess a distinctive streak at the base of the mandible (Short 1967). According to Short (1967), the broad black crown cap, trimmed by a thinner black line on the crown, is ancestral to grouse such as Lagopus, and all crown markings of Centrocercus and Dendragapus species are remnants of the full crown cap and trimmings of ancestral forms. This suggests that there is an evolutionary relationship between the head markings within the Tetraoninae.

What do we know about francolin and spurfowl chick plumage relationships?

Hall (1963) placed all spurfowls (occurring in Africa only)and francolins (occurring in Asia and Africa) under one genus, Francolinus, and described differences between chicks based on chick plumage. She divided the “monophyletic francolins” into eight taxonomic groups and mentioned that the chicks of the Bare-throated, Montane, Scaly, Vermiculated (all in the genus Pternistis),Striated (Dendroperdix) and Spotted (Francolinus)groups have a single conspicuous brown longitudinal band (with side lines) on the back through the crown(the crown cap) and a solid blackish stripe from the bill through the eye to the neck (extended eye stripe). The Red-winged (Scleroptila) group also have a variegated back, but with a narrower longitudinal band, and a narrower and darker crown patch than the spurfowls, with eight alternate black and white stripes on the sides of the crown, through and under the eyes.

Within the Red-tailed (Peliperdix) group, the female P. coqui chick is similar to the chicks of the Red-winged group in that it has a crown patch with lateral dark brown and buffy coloured stripes arranged across the back through the crown and stripes under the eyes. Conversely, the male P. coqui chick displays a similar pattern to the basic patterns displayed by the chicks of the Barethroated, Montane, Scaly, Vermiculated, Striated and Spotted groups, namely a broad crown cap and a single conspicuous rufous longitudinal band across the back through the crown cap and a solid blackish eye line. This difference between male and female P. coqui constitutes sexual dimorphism at the chick stage (Hall 1963).

In the Red-winged group the striped facial pattern (disruptive plumage) becomes a permanent cryptic feature in the adults, but in spurfowls the facial eye stripes often disappear, resulting in a plain colour in many species(Little and Crowe 2011). Although Hall (1963) mentioned that some variation was evident among sub-species of F. francolinus and F. shelleyi, she did not elaborate on this. Overall, she concluded that small variations existed among species and groups.

Francolins and spurfowls are not monophyletic

Crowe et al. (1992, 2006) reviewed the monophyletic genus “Francolinus” proposed by Hall and found that it represents two distant clades split amongfive genera:spurfowls (Pternistis) and francolins (Francolinus, Scleroptila, Peliperdix and Dendroperdix). In other words,based on DNA and morpho-behavioural studies, francolins and spurfowls do not share a common evolutionary path (Milstein and Wolff 1987; Crowe and Little 2004;Crowe et al. 2006). Bloomer and Crowe (1998) included the genetic material of several species, and Mandiwana-Neudani et al. (2014) analysed the genetic material (in conjunction with vocalisations) of 40 species (18 francolins and 22 spurfowls) to demonstrate the dichotomy between spurfowls and francolins. Mandiwana-Neudani et al. (2011) described the syringeal features that separated the two groups. However, none of these authors assessed chick plumage in any depth.

The Asian francolins have a rufous central longitudinal band on the back and crown, like G. gallus. The blotches behind the eye (as in F. gularis) are like some Tetraonidae spp., being especially evident in D. obscurus pallides and L. lagopus. Dendroperdix sephaena has all the characteristics of the G. gallus chick. However, the single diagnostic isolated spot on the neck is similar to a spot recorded in the same area of the neck of F. gularis and F. francolinus(Fig. 5). Peliperdix coqui chicks also have the characteristics of G. gallus, but the female chick’s striped facial parts are similar to those of the N. meleagris chick. Scleroptila gutturalis falls in the same category as the female P. coqui chicks. It has a crown and back feather pattern like G. gallus, but the facial parts resemble N. meleagris chicks.

Methods

We conducted phylogenetic analyses with common set of character states for francolin and spurfowl chicks in conjunction with a relatively large outgroup (Additionalfile 1: Fig. S1). A wide selection of outgroup species was important to determine the role that plesiomorphic character states play during the evolution of colours and patterns as suggested by existing literature (Short 1967; Jehl 1971) (Table 1).

Representative species from the groups described by Hall (1963) were examined, totalling 11 francolin species and 16 spurfowl species. All the chicks of francolins and spurfowls as well as a wide outgroup in the collections of the Natural History Museum at Tring and the American Museum of Natural History in New York were photographed to capture the dorsal and lateral characteristics of the head and the dorsal characteristics of back plumage (Milstein and Wolff 1987, Table 1). This data set was augmented with photographs from searches on the internet, drawings in reference books and unpublished photographs and sketches by the author (e.g. Milstein and Wolff 1987; Little and Crowe 2011; Little 2016). A photograph of a pair of Pternistis hartlaubi chicks was provided by Joris Komen. A day-old P. adspersus and P.natalensis hybrid chick was photographed by the author(JHvN unpublished). Although it was not possible to age the chick skins (age of chicks in days), spurfowl and African francolin chicks revealed head and facial markings and longitudinal bands on their backs which were generally consistent with the description of downy chick plumage, separating them from juveniles, in the literature(Crowe et al. 1986; Little 2016). Four Pternistis swainsonii chicks raised by a free ranging bantam Gallus domesticus hen on a farm in South Africa retained these markings for at least 5 weeks (JHvN unpublished). However,a study conducted by Little and Crowe (1992) showed that the back feathers of the Scleroptila afra chick disappears after about 14 days. Since all Scleroptila spp. had clear back bands, it is accepted that all Scleroptila spp.assessed were less than 14 days old. The plain state, or plain brownish or reddish chick plumage, was assumed as the ancestral root for the phylogenetic trees. In keeping with this, the Maleo (Macrocephalon maleo) is regarded as the oldest taxon among Galliformes; the chick possesses plain colours (not cryptic) (Crowe et al. 2006). The elected character states and the scores (0–2) allocated to 8 characters (and 25 states) are shown in Table 3. Figure 1 shows the parts (characteristics) of the head and back plumage that were used to compile the character matrix in Table 3. The taxa in Table 3 follow the nomenclature presented by Crowe et al. (2006), Little and Crowe (2011),and Mandiwana-Neudani et al. (2014) except the specific epithets which follow Gill and Donsker (2017). The mitochondrial Cytochrome b (mt Cytb) sequences including those of francolins, spurfowls and the selected outgroups were sourced from GenBank (Table 2).

Phylogenetic analyses—chick plumage and molecular characters

Parsimony analysis was performed on both chick plumage and Cytb characters in Windows-based PAUP* ver. 4.0b10 (Swofford 2002) with the following tree search parameters considered: full heuristic search with all characters unordered and with equal weight, starting tree(s) obtained via stepwise addition;tree-bisection-reconnection branch-swapping option was in effect, and 1000 random additions of taxa (Maddison 1991). One tree was held at each step during stepwise addition, with branches collapsed (creating polytomies) if the maximum branch-length was zero. Multiple, equally parsimonious cladograms were recovered and strict consensus cladograms were constructed. Bootstrap resampling (Felsenstein 1985) with 1000 pseudoreplicates andfive random additions of taxa per bootstrap pseudoreplicate was used to determine the degree of support for each node. Both the natal downy and Cytb phylogenies were rooted on M. maleo. Three measures of phylogenetic signal were gained by estimating consistency index(CI), which measures how well the data and the treefit with each other, that is, the consistency of the tree to the dataset; the retention index (RI), which measures how well synapomorphies explain the tree; and the homoplasy index (HI), which measures the amount of homoplasy observed on a tree relative to the maximum amount of homoplasy that could theoretically exist. These metrics reveal the amount of phylogenetic signals inherent in both character matrices.

Ancestral state reconstruction

The ancestral state reconstructions were made on the existing mt Cytb phylogeny for eight natal downy characters (Table 3) using Parsimony approach implemented in Mesquite ver. 3.2 (Maddison and Maddison 2017).All character states were unordered and the analysis employed the Markov k-state 1 parameter (mk1) model(Lewis 2001) where any particular change from one state to the other is equally probable.

Results

Broad groups of chick plumage

Author details

Plain: The darkish chicks of megapodes (Megapodiidae) are plain dorsally without stripes. The chick plumage of guineafowl species that live in the open understory of forests, such as Guttera pucherani and Agelastes meleagrides, are vaguely striped (rather blotchy) compared with the prominent striped patterns of guineafowl species that raise their chicks in grass, such as Numida meleagris and Acryllium vulturinum (Madge and McGowan 2002)(Fig. 2). The chick plumage of the forest-living A. niger is mainly dark rufous and black above (Little 2016).

Semi-striped: Grouse (Tetraonidae), Gallus gallus and Phasianus chicks have a variegated pattern with a central longitudinal band that proceeds through the crown to the forehead (Fig. 3). The rufous crown stripe often forms a single broad crown cap, with thinner trimmings on either side of the crown cap.

Striped: These chicks possess multiple stripes on the sides of their faces and are striped longitudinally across the crown right to the nares. Their back patterns are also variegated, but with a prominent central longitudinal band. Numida meleagris and A. vulturinum are typical examples of striped chicks (Fig. 2C, D). Coturnix coturnix chicks also display multiple stripes. The well-defined thin stripes across the crown and on the sides of the face of N. meleagris compared with the weakly defined stripes and blotches (fuidity) of A. meleagrides and G. pucherani suggest that the well-defined stripes emerged from a darkish plainer ancestor (see A. niger), with its black plain dorsal chick plumage. The latter three guineafowl species live in forests.

Chick plumage of spurfowl chicks

The 15 spurfowl chicks for which data were captured show a marked degree of similarity in terms of the selected character states (Table 3, Additionalfile 1: Table S1). The chick plumage of spurfowl chicks is regarded as semi-striped, since they lack the multiple disruptive stripes on the face and crown. The chick plumage of a hybrid P. adspersus and P. natalensis chick did not vary from this typical pattern. The back bears a broad, dark brown (rufous) central longitudinal band fanked by thinner and lighter-coloured stripes (otherwise known as a variegated back pattern). The crown has a broad rufous crown cap which is often trimmed by a thin,darker stripe on the edges. This crown cap is essentially an extension of the longitudinal band across the back.The face has an eye stripe which extends for a short distance in front of the eye (it often resembles a blotch in front of the eye). Some species, such as P. icterorynchus, P. nobilis and P. erckelii, generally have a reddish appearance, but their patterns (structurally) remain like the rest. P. hartlaubi is like the rest, with limited facial stripes and a variegated back pattern with a central longitudinal band on the back.

Chicks from two species showed prominent diagnostic features. Pternistis nobilis has a bold, thicker eye stripe,which is more prominent than the rest of the spurfowls,with a relatively thick frontal eye stripe. Pternistis erckelii has a solid, longish jaw stripe which in other species is faint or absent (Fig. 4).

利用灰色预测不能把原始观测数据作为一个简简单单的随机过程，而是把它看作一个随时间一起演变的灰色过程。通过对原始观测数据序列的处理加工，利用一次或者多次的累加累减使事物的灰色量达到白化的程度，突显事物内部的规律性，从而建立相应的变形预测微分方程，预测出事物的发展趋势。对于灰色模型来说，它所采用的数据既可以是线性的数据，也可以是非线性的数据，数据具有灵活性［1］。

Spurfowl chicks differ from their closest evolutionary relatives. Both the spurfowl chicks and Coturnix coturnix chicks are semi-striped on their heads but Alectoris chukar chicks possess stripes. Dorsally, only spurfowl chicks have a broad crown cap compared to scattered or banded crowns in the other two species (Table 3).

Chick plumage of francolin chicks

Data were captured for 11 species representing all four genera (Table 3).

Asiatic francolins: F. pondicerianus, F. francolinus and F.gularis are semi-striped. Like the spurfowls, they possess a rufous crown cap. The sides of their faces often have a few blotch-like blackish markings (probably remnants of stripes from an ancestor that was fully striped) behind the eye and on the neck. Like spurfowls, they also have a longitudinal band on the back with thinner, lighter-colouredlateral stripes (in other words, variegated back markings).Interestingly, only some F. pondicerianus chicks have a striped crown, showing intra-specific variability.

Dendroperdix sephaena is semi-striped, and shows resemblance to the spurfowls and Asiatic francolins. It has a rufous, crown cap with a solid eye line behind the eye, stretching down along the neck which is also typical for spurfowl chicks. It has a distinctive round dark spot on the neck, which it shares with F. francolinus and F. gularis (Fig. 5). The back also has a broad, dark brown central longitudinal band fanked with thinner,lighter-coloured stripes (in other words, variegated back markings) similar to spurfowl chicks.

施泰特的话中之意是让她考虑一下自己的牙齿，老妇人有所顾忌地说：“我如果买了黄桃罐头，你岂不是要少赚一些钱？”

The male P. coqui chick is semi-striped, with a rufous crown cap and a solid eye stripe behind the eye, a broad,dark brown longitudinal band on the back fanked by lighter lateral stripes reminiscent of spurfowl chicks(Fig. 6A, B). A female chick collected at Shangabue (Zimbabwe) shows a typical spurfowl crown and back, but the sides of the face have multiple stripes, which is typical of Scleroptila spp. (Fig. 6C). A similar female chick specimen is kept in the Ditsong National Museum of Natural History in South Africa. The distinction between female and male chicks is also evident at the juvenile stage, when the male has a clean buffy coloured face and the female multiple facial stripes; this is eventually carried through to adulthood. The male chicks resemble spurfowl chicks and are more closely related to the Asiatic chicks than to the striped chicks of Scleroptila spp.

Scleroptila gutturalis has a broad rufous crown cap,with multiple stripes on the face and a broad central longitudinal band on the back with thinner, light-coloured lateral stripes. Scleroptila levaillantii, S. shelleyi and S.afra have striped heads (instead of a well-defined crown cap) and facial markings, and the head colours are distinctively white and black, revealing a chevron pattern.

The Asiatic francolins, D. sephaena and the male P.coqui chick possess broad crown caps and semi-striped faces which are like their closest evolutionary relative G.gallus. Scleroptila species do not share these characteristics with G. gallus.

Chick plumage and Cytb phylogeny

The natal downy (Table 3) and Cytb data sets consisted of eight and 1,143 characters, consisting of eight and 430 parsimony informative characters, respectively. The matrices had a CI of 0.405, RI of 0.734 and HI of 0.595 whereas Cytb yielded a CI of 0.50, RI (of 0.521) and HI of 0.5, respectively. The Cytb phylogeny confirms the francolin-spurfowl dichotomy (Fig. 7) with francolins, spurfowls,grouse and allies (Tetraoninae) all in the family Phasianidae. Numididae is also recovered. The number of taxa included in this analysis had to match those for which the natal downy characters were available and this possibly could have resulted in the topology and nodal support that is slightly different from that in Crowe et al. (2006).The Asian/African linking lineage consisting of F. francolinus pondicerianus, F. gularis and D. sephana was recovered even though the branch from which these species arose is unresolved. The natal downy phylogeny (Fig. 8)shows poor phylogenetic structure which is largely polytomous and therefore no inferences can be made.

Symplesiomorphy underlying spurfowls

Ancestral characters form the “building blocks” that shape the colours and patterns of downy chicks. The broad longitudinal central rufous longitudinal band on the back and crown of spurfowl chicks (variegated) are also evident in G. gallus and grouse. The solid eye line of spurfowls, with the rest of the face often clean, is also evident in G. gallus, and is particularly similar between P.nobilis and G. gallus (see Figs. 3, 4). The prominent jaw stripe of P. erckelii (and to a lesser extent by other spurfowls) is also evident in Coturnix coturnix, N. meleagris and V. acryllium chicks.

Symplesiomorphy underlying francolins

We assessed the chick plumage characters of chicks of traditionally recognised spurfowl and francolin species, in conjunction with a broad selection of outgroup species, to (1) describe the chick plumage of spurfowls and francolins; (2) phylogenetically confirm or dismiss the dichotomy between spurfowls and francolins; (3)investigate the patterns of evolution of chick plumage characters which are diagnostic traced on a molecular phylogeny and (4) evaluate habitat matching of spurfowl and francolin chicks.

The chicks of S. afra, S. lavaillantii and S. shelleyi are similar to the striped pattern seen in N. meleagris, A.vulturinum and Coturnix coturnix chicks. However, the black lateral facial patterns of the latter three francolins stand out and could be related to the black facial parts of an Asiatic ancestor such as F. gularis. This is unlike S.gutturalis, which does not fall in this category and does not have black-coloured stripes. The plain colours of P.lathami chicks resemble the plain colours found in P.pucherani, A. meleagrides and A. niger.

Correlations between micro‑habitat and chick plumage character states

Micro-habitat refers to the immediate environment of the bird within a broader landscape. Generally, species clades live in uniform micro-habitats for most of the year (Fig. 9). Some, such as N. meleagris and A. vulturinum live out in the open but breed in closed habitat, in other words in grass. Peliperdix lathami is an exception as it lives in forests but its close relatives, P. coqui and P.albogularis move into open micro-habitats (Fig. 9).

Most species that live in open micro-habitats (Peliperdix, Dendroperdix, Francolinus spp., Pternistis spp.,L. lagopus, T. cupido and G. gallus) are semi-striped compared to Scleroptila spp., C. coturnix, N. meleagris and A. vulturinum that live in closed micro-habitats and are all multi-striped (more cryptic) (Additional file 1: Table S2). Most species that live in open microhabitats possess rufous variegated colour combinations but species that live in closed micro-habitats have dark reddish and rufous colour combinations.Most forest species possess reddish colours (Additional file 1: Table S2).

Generally, species that live in open micro-habitats possess less cryptic markings than species that live in closed micro-habitats or forests. Coturnix coturnix is closely related to Pternistis spp. but unlike Pternistis spp. they live in grasslands and therefore possess cryptic multiple stripes over their faces and crowns like the chicks of Scleroptila spp. to which they are not as closely related phylogenetically (Fig. 7).

我的画：流淌的水波纹是汇合？是守望？生命侵透在湿润的大气中彼此言说。我尝试用符号来串联、焕发出某种背后的思虑。

Most species that live in open micro-habitats possess a distinct eye band and comparatively most species that live in closed micro-habitats possess multiple facial stripes. Most forest species possess spots on the sides of their faces. Most species that live in open micro-habitats do not have jaw stripes whereas species that live in closed micro-habitats have partly or fully developed jaw stripes.Forest species do not possess any markings on the sides of their faces (Additionalfile 1: Table S2).

Dendroperdix sephaena and F. gularis have a clear isolated dark spot on the side of the neck, a discontinued break between the eye stripe and the neck stripe, a faint jaw stripe and an eye liner which supports the view of Mandiwana-Neudani et al. (2014) that D. sephaena phylogenetically resembles the Asiatic francolins and might be considered a linking form between the Asiatic and African francolins (Fig. 5, Table 3).

Most species that live in open micro-habitats possess broad stripes on their backs and species that live in closed micro-habitats possess thin and broad stripes. Comparatively forest species possess both but do not show any preference for any pattern (Additionalfile 1: Table S2).

Most species that live in open habitats possess no nose spots and for species that live in closed and forest habitats no preference was revealed. They both have spots or no spots. Generally, species of closed micro-habitats are more cryptic than species in open micro-habitats (Additionalfile 1: Table S2).

Ancestral state reconstruction

Disruptive markings

The ancestral state is inconclusive that is, the ancestor could have either been semi-striped or plain (with no markings). It seems semi-stripe is a plesiomorphic state among francolins and spurfowls with independent evolution of stripes in the grassland Scleroptila lineage, the spurfowls’ closest phylogenetic relative (Coturnix coturnix) and twice within Numididae and the sub-family Tetraoninae (Additionalfile 1: Fig. S2). There were two reversions to plain plumage in P. lathami and Tetrao urogallus.

Colour combinations of head

Generally, the ancestral states are inconclusive with the probability of all states (blackish brown, blackish, rufous variegated) having been present in the earlier ancestor.The colours of the head of M. maleo is plain black-brownish. This ancestral state together with the two diverged colour forms, rufous variegated and blackish colours, are found among the chicks of both Pternistis and Scleroptila spp. However, the head is consistently rufous variegated among Francolinus spp. (Additionalfile 1: Fig. S3).

Dorsal patterns

Generally, the ancestral states are inconclusive with the probability of all states (unchanged pail, banded, scattered, capped) having been present in the earlier ancestor. In M. maleo the dark head forms a continuous dark unchanged colour with the rest of the head and facial parts. The plain dorsal crown of M. maleo diverged into a well-defined capped crown found among Pternistis and Francolinus spp. but with a longitudinally banded pattern across the crown of Scleroptila species (Additionalfile 1:Fig. S4).

Side view patterns

The ancestral state is inconclusive, that is the ancestor may have had either plain or eye band/stripe. Francolins and spurfowls have eye band with independent evolution of multiple patterns in the grassland Scleroptila lineage and spots in the grouse and allies, Lagopus/Phasianus(Tetraoninae) lineage with an inconclusive ancestral state in members of Numididae. Unlike Scleroptila spp., the eye-stripes also evolved in Francolinus species (Additionalfile 1: Fig. S5).

Jaw stripe

With regard to this character, the ancestral state is plain(no distinguishable jaw stripe). Even though most of the deeper node states are inconclusive, the francolins and spurfowls generally have a plain state, the independent evolution of full (jaw stripe) occurred six times (thrice among francolins and once among spurfowls). The state“partly” stripe independently evolved one among francolins and twice among spurfowls. The grouse and allies is the lineage where the ancestral plain state persisted as well as in the linking lineage between Asian and African francolins (Additionalfile 1: Fig. S6).

Back colours

In summary, the following evolutionary pathways of Pternistis spp., Francolinus spp. and Scleroptila spp.might have played out: plain facial colours (see M. maleo)was followed by limited stripes, forming an eye-stripe in ancestral Phasianidae and was particularly deposited throughout Pternistis spp., Francolinus spp. and Peliperdix spp., followed by the emergence of some pigmentation in the form of multiple dark facial stripes in Scleroptila spp. In other words, when birds moved into grass, facial stripes became prominent. The scattered spots in grouse and allies (Tetraoninae) might be a breakdown of eye-stripes (divergence of eye stripes) since these spots are in that area of the neck towards where the eye stripe extends. However, from a dorsal point of view all variants (full pigmentation, cap and banded) were present among earlier ancestors of spurfowls and francolins. The facial sides of gamebird chicks (around eyes) are important for survival. The chick invariably uses his head as look-out point and will therefore adapt its morphological appearance to match the micro-habitat for camoufage and still allow maximum vigilance, while the rest of body is lower and often hidden. It is understandable that ancestral states diverged into multiple forms (that is the inconclusive linking of the ancestral states to Pternistis and Scleroptila spp.) because this is ascribed to habitat matching. Despite that the symplesiomorphic states hinders the use of downy characters to its full potential for the purpose of classification, this study revealed the evolutionary patterns of various characters and also how they associate with habitat. This had never been explored.

Back patterns

Generally, the ancestral states are inconclusive with the probability of all states (plain, narrow, broad) having been present in the earlier ancestors. The M. maleo chick is black-brownish plain. This state diverged into two clear patterns. Most Pternistis spp. possesses a broad central back band fanked by two thinner but lighter coloured stripes (dilution of pigmentation) on either side. All Scleroptila spp. have markedly narrower back stripes which evolved independently and this is observed in P. rufopictus. Peliperdix lathami experienced a throw-back of plain colours on their backs as found in M. maleo. The re-appearance of plain back is also observed in T. cupido and A. meleagrides (Additionalfile 1: Fig. S8).

Nose spot

The ancestral state is plain (no distinguishable nose spot).Macrocephalon maleo does not possess a nose spot since the nose and crown consist of a continuous black-brownish plain colour. The absence of a nose spot prevails throughout the francolin and spurfowl lineage (pleisiomorphic state) with independent evolution of nose spots within the Red-winged grassland Scleroptila subclade,among spurfowls that is, among the Montane, Vermiculated species as well as in the most basal spurfowl species P. hartlaubi. Importantly, the plain state of ancestors such as M. maleo is due to full pigmentation while the plain state among francolin and spurfowl is due to the dilution of pigmentation (Additionalfile 1: Fig. S9).

Discussion

Phylogeny

From a cladistics point of view, the topology of the natal downy phylogeny of francolins and spurfowls is largely unsatisfactory or polytomous. This could be due to the high degree of symplesiomorphy within Phasianidae (Johnsgard 2008) (Fig. 8). It was therefore not possible to delimit species satisfactorily based on plumage characters. These results are incongruent with the spurfowland francolin phylogeny built on DNA and adult morpho-behavioural characters wherein resolved species relationships were determined with a high nodal support (Mandiwana-Neudani et al. 2014). The latter refutes Hall’s (1963) hypothesis and confirm that spurfowls(Pternistis) and francolins (Francolinus, Dendroperdix,Peliperdix and Scleroptila) do not form a monophyletic group (Crowe et al. 2006). Thus, the spurfowl and francolin phylogeny based on natal chick plumage characters does not support this dichotomy.

Inter‑specific comparisons

The back colours of species that live in open microhabitats are rufous and most species that live in closed micro-habitats also possess rufous colours. Comparatively forest species possess dark reddish colours on their backs (Additionalfile 1: Table S2).

Female P. coqui chicks have the disruptive eye stripes seen in S. gutturalis, and the male has limited facial markings as seen in the spurfowl chicks, which places this species between the two clades based on chick plumage.Hartlaub’s Spurfowl P. hartlaubi is regarded as the most basal species of all extant spurfowls, but has a mating system and vocalisations which differ substantially from all other spurfowls (Komen 1987; Van Niekerk 2013). There is some debate as to whether this species belongs to the genus Pternistis, but based on chick plumage it is similar to all spurfowls (J. Komen in litt). It was expected that the spurfowl chicks of P. ahantensis, P. squamatus and P. griseostriatus that live on the fringes of forests would have attained forest-like character states such as a dark, reddish, rufous, plain dorsal plumage or at the least vague blotches, instead of variegated stripes fusing into a darker background plumage colour. In other words, it was expected that the forest species would resemble other forest-living birds such as A. meleagrides, G. pucherani and P. lathami and the forest-dwelling tinamous chicks(Jehl 1971), all of which have plumage patterns which are not contrasting and generally darker. Instead they are very like all the other savannah-living spurfowls. Lack of forest-living characteristics among the spurfowls that live on the fringes of forests could be ascribed to a relatively recent speciation event, leaving these taxa in a state of fux, engaged infinding and refining habitat niches which may, in evolutionary time scales, imprint on chick plumage patterns and colours (Milstein and Wollf 1987). This lack of deviation from unity is supported by the polytomous structures in the cladograms (Fig. 8).

在酸醚比3:1、小单体总量和丙烯酸用量不变、引发剂用量为1%、链转移剂用量为0.9%，考察了疏水单体醋酸乙烯酯(VAc)用量对减水剂性能的影响，结果见图3。

规范的临床案例包括主诉、现病史、既往史、实验室检查、临床处理过程5个部分，将大纲要求的各知识点整合起来，同时整合学生所学专业、人文、社会、伦理等知识，教师引导学生剖析案例，将习得的纵向知识进行横向联系、融会贯通，学会解决临床实际问题，既巩固已学知识，又提高学生综合素质，为学生终身发展奠定基础。

No character captured from spurfowl or francolin skins could be described as “derived” (not observed in any ancestor), which confirms the strong role that symplesiomorphy plays in natal chick plumage within Phasianidae, as was demonstrated within grouse (Short 1967)and which also seems applicable to other families such as that of the tinamous (Bertelli et al. 2002). In fact, at the outset it was demonstrated that spurfowls and francolins could each be divided into three broad plumage categories found in the wider Phasianidae family, namely plain,striped and semi-striped chicks, which confirms the role played by symplesiomorphy.

二战后独立的东南亚民族国家中兴起的民族主义意识也对海外华人跨界企业家的价值和地位发出挑战和质疑。海外华人跨界企业家创业发展的起起伏伏与华人企业家自身的移民身份、少数民族地位、在华人社团中的领导角色和与所在地权力精英的关系息息相关。华人移民企业家的地位取决于他们在所在国家和地区的网络构建能力、创业实践成就和在当地的社会政治文化参与程度。华人少数民族企业家的形成与他们在东南亚、北美、欧洲等居住地区的政治、经济、社会发展历史进程紧密地连在一起。当他们回到中国之时又卷入了中国改革发展的进程当中。

日前，我国满载6.15万吨LNG的“Gaslog Singapore”号顺利抵达中国海油粤东LNG接收站。这意味着我国进口LNG窗口一站通首期试点产品进入船货交割阶段，标志着我国实现了LNG接收站向第三方公平开放。进口LNG窗口一站通产品是中国海油气电集团与上海石油天然气交易中心共同推出的国内首个对外开放的窗口期产品。该产品的试点于9月20日在交易中心平台完成线上交易，由振华石油和胜通能源组成的联合体通过公开竞价购得。

Field identification

It was possible to separate subsets according to plumage characteristics. Scleroptila show strong signs of disruptive lateral facial patterns. Conversely, the spurfowls have limited facial stripes, but a broader longitudinal band running from the crown down the neck and through the centre of the back. These differences have also been reported in the literature (Little 2016). The Asian francolins have a broad longitudinal band on the back like the spurfowls, but have black spots or blotches (speckled remains of stripes) just behind or below the eye, which is more in accordance with the spurfowls than for Scleroptila spp. Asian francolins are similar to D. sephaena,which has a single round dark spot just below the ear.

占有是指在现实生活中对某一事物的支配控制，也是其占用人有权对其具有的价值进行利用和管理。农民拥有土地产权的条件是拥有土地，只有这样，农民才能拥有土地产权的基本权利，即土地收益权和使用权。但是在我国，土地是属于国家的，相对个人而言，农民可以通过土地承包制获得一定年限的土地使用权、收益权等。农民目前对土地还没有占有权[1]。

Pternistis erckelii is similar to the rest of the spurfowls,but has a distinct jaw line which is only partly visible on the jaws of a few other spurfowls. The plain back plumage of Peliperdix lathami (formerly F. lathami; Hall 1963; Madge and McGowan 2002) is at variance with the spurfowls and francolins that have dorsal stripes. These plain dorsal features on the back could be an adaptation for life in forest conditions to break the bird’s outline in shade, whereas semi-striped or striped plumage might be detected by predators under these conditions (Stevens and Merilaita 2008). A more complete set of francolin and spurfowl skins than what is currently available may reveal useful diagnostic features for individual species, but it is doubtful whether it would reveal derived traits that would assist in describing speciation based on chick plumage.

Habitat matching

Most species that live in open micro-habitats possess rufous caps on their crowns compared to all species living in closed micro-habitats that possess banded crowns.Forest species possess caps as well (Additionalfile 1:Table S2).

Scleroptila species. are not closely related to Numida meleagris or Acryllium vulturinum but as the latter species raise their chicks in grass like Scleroptila spp., they all possess cryptic stripes like Scleroptila spp. Peliperdix lathami lives in forests and therefore does not possess an eye stripe like Peliperdix coqui, Peliperdix algularis, Francolinus spp. and Pternistis spp. that live in open microhabitats. Peliperdix spp., Dendroperdix sephaena and Francolinus spp. are more closely related to Scleroptila spp. but unlike Scleroptila spp. they live in open macrohabitats and therefore unlike Scleroptila spp., they do not possess multiple stripes across their faces and crowns,but instead possess a single wide eye stripe like Pternistis spp. which live in open micro-habitats. Gallus gallus are more closely related to Scleroptila spp. but their chicks look like the chicks of Pternistis spp. since they also live in open micro-habitats. It is therefore concluded that habitat matching is a likely driving force that determines the natal down of francolin and spurfowl chicks.

本研究结果表明,观察组社区糖尿病患者治疗总有效率98.00%,相比对照组更具有优势(P<0.05)。观察组社区糖尿病患者不良反应发生率4.00%,低于对照组,2组间相比较,P<0.05。

Does the chick plumage of spurfowl and francolin chicks have any significance for camoufage? Why don’t these chicks simply have plain colours like A. meleagrides or Peliperdix lathami that live under the canopy of forests? The limited facial stripes of spurfowl chicks probably assist the chicks to blend in with a bushier type habitat with limited ground cover such as grass. Should these chicks possess multiple stripes it would be conspicuous under these open conditions as it would define the outline of the head (Merilaita and Lind 2005; Stevens and Merilaita 2008). On the other hand, the multiple facial stripes of S. afra, S. levaillantii and S. levaillantoides chicks possibly break the outline of the shape of the chick’s head in grass cover. A non-striped head will be more detectable in grass than is a striped head (Merilaita and Lind 2005). In both francolins and spurfowls, the variegated back stripes probably provide countershading,making it diffcult for a predator to distinguish the shape of the chick from its physical surroundings (Caro 2014).

In summary, although symplesiomorphy is a hereditary explanation for downy colours and patterns, the traits relevant for habitat matching are combined in a manner which is determined (adaptation) by natural selection(Frost 1975; Paxton 2009; Peichel 2014).

Evolution of downy patterns

Macrocephalon maleo is the most basal outgroup species to spurfowl and francolin within the Galliformes.Its downy chick plumage is basically uniformly blackbrownish. The divergence of other downstream variants is basically a dilution of darker pigment (Mills and Patterson 2009). While speciation continued dark plain colours became limited but patterned on certain body parts of descendants, particularly on the crown, face and back.A. meleagrides and Guterra pucherani are proximate descendants to M. maleo and here it is evident that the uniformly dark plumage of downy chicks possesses whitish (diluted) blotches, but not yet particularly sharply defined features (e.g. striped).

Further down the evolutionary pathway two overriding morphological forms emerged with less black-brownish pigmentation in certain areas revealing dark stripes on their heads and back that are symmetrical and orderly arranged on a buff-coloured (diluted) background. These patterns are observed among many Phasianidae but particularly among Pternistis spp. where it emerged as semistripes (fewer stripes) compared to Scleroptila spp. where the remaining rufous stripes emerged as multiple stripes,giving the bird a disruptive chick plumage.

糖尿病患者服药依从性是决定其预后的重要影响因素。老年患者的依从性不仅与其年龄、文化程度和经济状况有关，还受到患者对疾病与治疗的知识、信念、心理状况以及患者所能得到的社会支持等因素的影响。而加强健康教育、改善家庭支持以及给予足够的心理支持是提高患者服药依从性的有效措施，并可通过这些措施将血糖控制在理想水平，延缓并发症的发生，延长寿命，提高生命质量。

The presence of nose spots happened when most darker pigmentation receded leaving two isolated spots (or short stripes) on either side of the base of the upper mandible(nares). Indeed, in some species such as Lagopus lagopus,Scleroptila afra and Numida meleagris there are continuous stripes from the nose across the orbit on both sides of the head showing that dilution emerged along stripes. The presence of jaw stripes and multiple stripes on the face is ascribed to the evolution of darker pigmentation in this area. An ancestral bird like M. maleo has uninterrupted dark plain colours all over the head but the darker pigmentation diluted on the lower facial areas, but leaving no distinguishable jaw stripe (Additionalfile 1: Fig. S1).

There are more stripes on the side of the face of Pternistis erckelii than in other spurfowls, such as having a distinct jaw stripe and a thin stripe above the orbit which is more bold and typical of Scleroptila spp. and female P.coqui. The jaw stripe is only faintly visible on some other spurfowls. Ostensibly, while this seems to be a logical course of evolutionary events, for instance that P. erckelii could link the spurfowls and Scleroptila spp., it is more likely that the ancestors of extant francolins and spurfowls moved back and forth between striped and semistriped chick plumage, which was probably determined by exposure to different habitats.

Conclusions

Generally, the ancestral states are inconclusive with the probability of all states (blackish-brown, blackish, rufous variegated) having been present in the earlier ancestor.The general back colour of the M. maleo chick is plain black-brownish. This ancestral character state plus a diverged rufous variegated pattern became characteristic among Pternistis spp. but Francolinus spp. only attained the rufous variegated patterns. Scleroptila spp. possess these variants but also include a blackish tone that is often revealed in the chicks of grouse and allies in the subfamily Tetraoninae. In short, all the ancestral states are consistently distributed across the tree. The exception if the independent evolution of a state shown by a white ball with black lines leading to a montane species P. castaneicollis and S. shelleyi which indicates that the observed state is undecided while the most parsimonious state is black brownish and rufous variegated, respectively (Additionalfile 1: Fig. S7).

Additionalfile

Additionalfile 1: Table S1. Characters, associated states and their scores as given in Table 3. Table S2: Counts of character states associated with species arranged under three micro habitats to demonstrate associa‑tion of species with habitat type. Figure S1: Facial, crown and dorsal feather patterns of the chick selected for this study. Figure S2: Ancestral state reconstruction for francolins and spurfowls based on Parsimony optimisation of disruptive markings traced on mitochondrial Cytochrome b phylogeny. Figure S3: Ancestral state reconstruction for francolins and spurfowls based on Parsimony optimisation of colour combination of head traced on mitochondrial Cytochrome b phylogeny. Figure S4:Ancestral state reconstruction for francolins and spurfowls based on Parsimony optimisation of dorsal patterns (crown) traced on mitochon‑drial Cytochrome b phylogeny. Figure S5: Ancestral state reconstruction for francolins and spurfowls based on Parsimony optimisation of side view patterns traced on mitochondrial Cytochrome b phylogeny. Figure S6: Ancestral state reconstruction for francolins and spurfowls based on Parsimony optimisation of jaw stripe traced on mitochondrial Cytochrome b phylogeny. Figure S7: Ancestral state reconstruction for francolins and spurfowls based on Parsimony optimisation of back colours traced on mitochondrial Cytochrome b phylogeny. Figure S8: Ancestral state reconstruction for francolins and spurfowls based on Parsimony optimisa‑tion of back patterns traced on mitochondrial Cytochrome b phylogeny.Figure S9: Ancestral state reconstruction for francolins and spurfowls based on Parsimony optimisation of nose spots traced on mitochondrial Cytochrome b phylogeny.

Authors’ contributions

JHvN produced the research design including conceptualisation, museum assessments and character analysis. TGMN conducted the phylogenetic analysis and the ancestral state reconstructions as well as the interpretations. Both authors have read and approved thefinal manuscript.

From the outgroup it was possible to select three broad groups of chick plumage colours and patterns by which to categorise the francolin and spurfowl chicks.

1 Department of Environmental Sciences, College of Agriculture and Environmental Science, University of South Africa, PO Box 392, Pretoria 0003, South Africa. 2 Department of Biodiversity, University of Limpopo, Private Bag X1106,Sovenga 0727, South Africa.

Acknowledgements

GB/T 20258.1―2007《1∶500 1∶1 000 1∶2 000基础地理信息要素数据字典》发布虽已超过5年，但在调研中了解到各单位尤其是城建部门才开始学习和使用该标准，也有部分单位根据该标准编制了自己内部的建库规范，他们希望保持其稳定性，不要做大的改动。因此，本次数据字典修订保持标准的稳定性，尽可能不做大的变化，修订的重点放在新增要素以及与图式、分类代码相矛盾的要素上。

We thank Petra van Niekerk for assisting with the selection and photography of museum skins and Melissa van Niekerk for drawing the chicks’ sketches.

Competing interests

The author declares that they have no competing interests.

Availability of data and materials

The datasets used and/or analysed during the current study are available from the corresponding author and are available for public access on GenBank.

Consent for publication

Not applicable.

Ethics approval and consent to participate

Not applicable.

Funding

Not applicable.

References

Berruti A. The AGRED guide to gamebird management in South Africa.Houghton: AGRED; 2011.

Bertelli S, Norberto PG, Golobof PA. A phylogeny of the tinamous (Aves:Palaeognathiformes) based on integumentary characters. Syst Biol.2002;51:959–79.

Bloomer P, Crowe TM. Francolin phylogenetics: molecular, morphobehavioral,and combined evidence. Mol Phylogenet Evol. 1998;9:236–54.

Caro T. Antipredator deception in terrestrial vertebrates. Curr Zool.2014;60:16–25.

Crowe TM, Little RM. Francolins, partridges and spurfowls: what’s in a name?Ostrich. 2004;75:199–203.

Crowe TM, Keith GS, Brown LH. Galliformes in birds of Africa, vol. ii. London:Academic Press; 1986.

Crowe TM, Harley EH, Jakutowicz MB, Komen J, Crowe AA. Phylogenetic,taxonomic and biogeographical implications of genetic, morphological,and behavioural variation in francolins (Phasianidae: Francolinus). Auk.1992;109:24–42.

Crowe TM, Bowie RCK, Bloomer P, Mandiwana TG, Hedderson TAJ, Randi E,Pereira SL, Wakeling J. Phylogenetics, biogeography and classification of, and character evolution in, gamebirds (Aves: Galliformes): Effects of character exclusion, data partitioning and missing data. Cladistics.2006;22:1–38.

Felsenstein J. Confidence limits on phylogenies: an approach using the Bootstrap. Evolution. 1985;39:783–91.

Frost PGH. The systematic position of the Madagascan Partridge Magaroperdix madagascariensis (Scopoli). Bull Br Ornithol Club. 1975;95:64–8.

Gill F, Donsker D, editors. IOC world bird list (v 7.2); 2017. https://doi.org/10.14344/ioc.ml.7.2.

Hall BP. The francolins, a study in speciation. Bull Br Mus. 1963;10:107–204.

Jehl JR. The colour patterns of downy young ratites and tinamous. Trans Soc Nat Hist San Diego. 1971;16:291–302.

Johnsgard PA. Molts and plumages. Grouse and quails of North America.Papers in the Biological Sciences. Lincoln: University of Nebraska; 2008.

Komen J. Preliminary observations of the social patterns, behaviour and vocalisation of Hartlaub’s Francolin. S Afr J Wildl Res Suppl. 1987;1:82–6.

Lewis PO. A likelihood approach to estimating phylogeny from discrete morphological character data. Syst Biol. 2001;50:913–25.

Little R. Terrestrial gamebirds & snipes of Africa. Johannesburg: Jacana Media;2016.

Little RM, Crowe TM. The use of morphometrics and development of plumage in estimating the growth patterns and age of greywing francolin Francolinus africanus. Ostrich. 1992;63:172–9.

Little R, Crowe T. Gamebirds of Southern Africa. Cape Town: Struik; 2011.

Livezey BC. A phylogenetic analysis and classification of recent dabbling ducks (tribe Anatini) based on comparative morphology. Auk.1991;108:471–507.

Maddison DR. The discovery and importance of multiple islands of most parsimonious trees. Syst Biol. 1991;40:315–28.

Maddison WP and Maddison DR. Mesquite: a modular system for 3.2. http://mesquiteproject.org. 2017. Accessed 11 Sept 2017.

Madge S, McGowan P. Pheasants, partridges and grouse. London: Christopher Helm; 2002.

Mandiwana-Neudani TG, Kopuchian C, Louw G, Crowe TM. A study of gross morphological and histological syringeal features of true francolins (Galliformes: Francolinus, Scleroptila, Peliperdix and Dendroperdix spp.) and spurfowls (Pternistis spp.) in a phylogenetic context. Ostrich. 2011;82:115–27.

Mandiwana-Neudani TG, Bowie RCK, Hausberger M, Henry LM, Crowe TM. Taxonomic and phylogenetic utility of variation in advertising calls of francolins and spurfowls (Galliformes: Phasianidae). Afr Zool. 2014;49:54–82.

Merilaita S, Lind J. Background-matching and disruptive coloration, and the evolution of cryptic coloration. Proc R Soc B. 2005;2005(272):665–70.

Mills MG, Patterson LB. Not just white and black: pigment pattern development and evolution in vertebrate. Semin Cell Dev Biol. 2009;20:72–81.

Milstein P le S, Wolff SW. The oversimplification of our francolins. S Afr J Wildl Res. 1987;1:58–65.

Paxton EH. The utility of plumage coloration for taxonomic and ecological studies. Open Ornithol J. 2009;2:17–23.

Peichel CL. Genetics of phenotypic evolution. In: Losos J, editor. Guide to evolution. Princeton: Princeton University Press; 2014. p. 452–7.

Short LL Jr. A review of the genera of grouse (Aves, Tetraoninae). American Museum novitates; No. 2289. New York: American Museum of Natural History; 1967.

Stevens M, Merilaita S. Animal camou fl age: current issues and new perspectives. Phil Trans R Soc B. 2008;364:423–7.

Swofford DL. PAUP*: analysis using parsimony. Version 4. Sunderland: Sinaeur Associates; 2002.

Van Niekerk JH. Vocal structure behaviour and partitioning of all 23 Pternistis spp. into homologous sound (and monophyletic) groups. Chin Birds.2013;4:210–31.

Vázquez DP, Gittleman JL. Biodiversity conservation: does phylogeny matter?Curr Biol. 1998;8:379–81.

Whetherbee DK. Natal plumages and downy pteryloses of passerine birds of North America. Bull Am Mus Nat Hist. 1957;113:343–436.