In North America and Eurasia, the most ancestral populations of species often occur in the southern portions of their ranges (Hewitt, 2000;Petit et al., 2003). These southern regions tend to be the most climatically stable and therefore served as refugia during the heights of ice-age glaciations. In contrast, northern regions were associated with dramatic climate changes, fueling an alternation of range expansions and range contractions or extinctions in many species (Hewitt, 2000;Hofreiter & Stewart, 2009). Populations that today are widespread across northern latitudes often trace their origins to relatively recent, late Pleistocene expansion events from

Vocal divergence within species often corresponds to morphological, environmental, and genetic differences between populations. Wolf howls are long-range signals that encode individual, group, and subspecies differences, yet the factors that may drive this variation are poorly understood. Furthermore, the taxonomic division within the Canis genus remains contended and additional data are required to clarify the position of the Himalayan, North African, and Indian wolves within Canis lupus. We recorded 451 howls from the 3 most basal wolf lineages—Himalayan C. lupus chanco—Himalayan haplotype, North African C. lupus lupaster, and Indian C. lupus pallipes wolves—and present a howl acoustic description within each clade. With an additional 619 howls from 7 Holarctic subspecies, we used a random forest classifier and principal component analysis on 9 acoustic parameters to assess whether Himalayan, North African, and Indian wolf howls exhibit acoustic differences compared to each other and Holarctic wolf howls. Generally, both the North African and Indian wolf howls exhibited high mean fundamental frequency (F0) and short duration compared to the Holarctic clade. In contrast, the Himalayan wolf howls typically had lower mean F0, unmodulated frequencies, and short howls compared to Holarctic wolf howls. The Himalayan and North African wolves had the most acoustically distinct howls and differed significantly from each other and to the Holarctic wolves. Along with the influence of body size and environmental differences, these results suggest that genetic divergence and/or geographic distance may play an important role in understanding howl variation across subspecies.

Natural and human-driven selection of a single noncoding body size variant in ancient and modern canids Highlights d An ancestral variant on IGF1 locus regulates body size in ancient and modern dogs d Variant alleles are associated with body size in dogs, wolves, and coyotes d The large body size-associated allele arose more than 53,000 years ago in wolves

The red wolf (Canis rufus) of the eastern US was driven to near‐extinction by colonial‐era persecution and habitat conversion, which facilitated coyote (C. latrans) range expansion and widespread hybridization with red wolves. The observation of some grey wolf (C. lupus) ancestry within red wolves sparked controversy over whether it was historically a subspecies of grey wolf with its predominant “coyote‐like” ancestry obtained from post‐colonial coyote hybridization (2‐species hypothesis) versus a distinct species closely related to the coyote that hybridized with grey wolf (3‐species hypothesis). We analysed mitogenomes sourced from before the 20th century bottleneck and coyote invasion, along with hundreds of modern amplicons, which led us to reject the 2‐species model and to investigate a broader phylogeographic 3‐species model suggested by the fossil record. Our findings broadly support this model, in which red wolves ranged the width of the American continent prior to arrival of the grey wolf to the mid‐continent 60–80 ka; red wolves subsequently disappeared from the mid‐continent, relegated to California and the eastern forests, which ushered in emergence of the coyote in their place (50–30 ka); by the early Holocene (12–10 ka), coyotes had expanded into California, where they admixed with and phenotypically replaced western red wolves in a process analogous to the 20th century coyote invasion of the eastern forests. Findings indicate that the red wolf pre‐dated not only European colonization but human, and possibly coyote, presence in North America. These findings highlight the urgency of expanding conservation efforts for the red wolf.

Vocal communication in social animals plays a crucial role in mate choice, maintaining social structure, and foraging strategy. The Indian grey wolf, among the least studied subspecies, is a social carnivore that lives in groups called packs and has many types of vocal communication. In this study, we characterise harmonic vocalisation types of the Indian wolf using howl survey responses and opportunistic recordings from captive and nine packs (each pack contains 2–9 individuals) of free-ranging Indian wolves. Using principal component analysis, hierarchical clustering, and discriminant function analysis, we found four distinct vocalisations using 270 recorded vocalisations (Average Silhouette width Si = 0.598) which include howls and howl-barks (N = 238), whimper (N = 2), social squeak (N = 28), and whine (N = 2). Although having a smaller body size compared to other wolf subspecies, Indian wolf howls have an average mean fundamental frequency of 422 Hz (±126), which is similar to other wolf subspecies. The whimper showed the highest frequency modulation (37.296±4.601) and the highest mean fundamental frequency (1708±524 Hz) compared to other call types. Less information is available on the third vocalisation type, i.e. ‘Social squeak’ or ‘talking’ (Mean fundamental frequency = 461±83 Hz), which is highly variable (coefficient of frequency variation = 18.778±3.587). Lastly, we identified the whine, which had a mean fundamental frequency of 906Hz (±242) and is similar to the Italian wolf (979±109 Hz). Our study’s characterisation of the Indian wolf’s harmonic vocal repertoire provides a first step in understanding the function and contextual use of vocalisations in this social mammal.

The gray wolf (Canis lupus; Linnaeus, 1758) is one of the most widespread terrestrial species and occurs in a variety of habitats. While well studied in North America and Europe, wolf populations in Asia are among the most evolutionarily distinct, endangered and data deficient. The Indian wolf (Canis lupus pallipes) is a subspecies of gray wolf that ranges from Southwest Asia to the Indian Subcontinent. Despite being categorized as “Endangered” in Pakistan, data on the gray wolf’s status, ecology and distribution are poorly understood. The current study investigates its genetic distinctiveness, distribution, feeding ecology and wolf livestock conflict in the Suleman Range, South Waziristan, Pakistan. We confirmed that the gray wolf is present in South Waziristan and is genetically similar to the wolves of Iran and Saudi Arabia based on their mtDNA D-loop haplotypes. The gray wolf was recorded at eight different sampling sites in the study area with elevational range between 1642 m to 2688 m. We estimated a population of 15 wolves, with a density of 0.62 individuals/km2 area surveyed. An analysis on scats revealed 52% contribution from livestock (with goats and sheep being the preferred prey) and 48% from wild prey. Biomass consumption showed gray wolf relied heavily on domestic prey (88%) during the summer season, resulting in human conflict with 28 wolves killed in response to livestock depredation during 2016–2017, requiring immediate conservation measures to save its remaining population.

PurposeTo investigate the relationship between phylogeny and amount of shade in a species’ habitat regarding the presence or absence of an iridal granula iridica (GI) in a large sample of Artiodactyl and Perissodactyl clades and using online resources.MethodsThe Comparative Ocular Pathology Laboratory of Wisconsin (COPLOW) archives were searched for glass slide material from Artiodactyl (even‐toed) and Perissodactyl (odd‐toed) ungulates. The slides were examined, and the presence or absence of the GI was noted. The phylogenetic tree of the ungulate species was inferred using TimeTree (http://www.timetree.org), and the habitat data are derived from Animal Diversity Web (https://animaldiversity.org/). We assessed the probability of the presence of GI occurring given the amount of shade in a species’ environment using phylogenetic logistic regression.ResultsForty‐eight artiodactyl species were able to be evaluated and tabulated. Nine perissodactyl species were able to be evaluated. The phylogenetic logistic regression showed that the probability of GI presence was lower in artiodactyl species that inhabited shaded environments (βshaded = −1.774). Arctiodacyl species inhabiting a nonshaded environment were slightly more probable to have the GI present (βnonshaded = 0.023), with species inhabitating ambiguously shaded environments having a high probability of GI presence (βambiguous = 2.214).ConclusionsOur results suggest that the GI may be a common morphological feature to shade the pupil in nonshaded environments, and, in its absence, increase the amount of light reaching the retina to improve vision in shaded environments for hooved mammals. Further research on the functional optics of the GI and studies that include additional ungulate species would further elucidate phylogenetic and ecological factors influencing the occurrence of GI in hooved mammals.

38 Vocalisation plays a critical role in social animals for conveying information on 39 foraging, reproductive, and social behaviours [1][2][3][4][5][6][7]. Characterising the vocal repertoire 40 of a species provides a base for understanding the behavioural significance of different 41 vocalisations and studying how vocal communication varies across the populations, 3 42 subspecies, and taxa [8][9][10]. The wolf (Canis lupus) is a social mammal and uses a 43 variety of vocalisations for communication. Being present throughout Eurasia and North 44 America, the wolf is one of the most widely distributed land mammals and occupies a 45 wide range of different habitat types [11]. The Indian wolf is among the smallest [12] 46 and one of the most evolutionarily distinct wolf subspecies, having diverged around 47 270,000 and 400,000 years ago based on mitochondrial DNA [13][14][15]. Studying the 48 vocal repertoire of the Indian wolf can aid in future studies on the function of different 49 vocal signals in Indian wolves and, more broadly, the variation in vocalisation across 50 subspecies and taxa within the Canis clade.51 The best-known wolf vocalisation -the howl -is a long-range harmonic call used for 52 territorial advertising and social cohesion [1,[16][17][18]. Recent studies have shown 21 53 different howl types across various canid subspecies based on quantitative similarity in 54 modulation pattern [8]. Along with howl, wolves also communicate using 7 to 12 other 55 harmonic calls, which is a clear pitch sound wave that possesses multiple integral 56 frequencies [19][20][21]. Many of these other harmonic vocalizations are short-ranged, and 57 due to difficulties in recording these calls, remain less studied compared to the wolf 58 howl [22]. These short-ranged calls are important for communicating passive or 59 aggressive behaviour among social canids [22][23][24].60 The whimper, whine and yelp are various calls for communicating passive and friendly 61 behaviour among wolves [18,23], whereas noisy calls, which don't have a clear pitch or 62 distinct frequency band in their spectrograms, communicate different levels of 63 aggression [18,23]. The whimper and whine vocalization is similar to a crying sound 64 with the whimper having a comparatively shorter duration than whine [18,25]. The 65 whine vocalization is mostly used for submissive behaviour whereas the whimper is 4 66 primarily used for greeting [18]. Yelp is a short and sharp cry and is used in submissive 67 behaviour with body contacts [18,25]. To communicate different levels of aggression 68 behaviors, wolves use noisy calls which consist of the growl, woof, and bark. Growl is a 69 laryngeal sound to show dominance in any interaction, whereas the woof vocalization is 70 a non-vocal sound cue (without involvement of vocal cords) used by adults for their 71 pups [18,25,26]. The bark is a short low pitched sound with rapid frequency modulation 72 and is used during aggressive defence [19,26], such as defending pups or defending a 73 food resource. Wo...