Primates

Many primates exhibit female philopatry and live in stable, female-bonded social groups. Permanent group fusions are rarely documented in these populations. We present a case study on a fusion of two social groups from a hybrid population of baboons (Papio cynocephalus x P. anubis) living in the Amboseli basin of Kenya. The fusion occurred following a period of increased human-induced mortality in one of the two social groups. After the fusion, females from the smaller group became the lowest ranking. We compared female behavior in the months following the fusion to the behavior of females in groups that had not fused and also compared pre- and post-fusion fitness outcomes. Following the fusion, the groups activity budget and patterns of agonistic interactions were typical for the study population. Females preferred familiar grooming partners for a short period following the fusion; however, after three months, patterns in female grooming were comparable to other groups, indicating rapid social integration. With the caveat that our sample size was limited, we observed no detectable fitness-related costs of group fusion in terms of birth rates or offspring survival, and adult female mortality was low following the fusion. These results demonstrate the flexibility of female baboons in navigating exposure to novel same-sex conspecifics despite a species-typic pattern of female philopatry. Based on this and previous examples of group fusions, we propose that group fusions may be most likely to occur when groups are too small to retain adult males, defend against predators, or compete with other groups.

In nature, the most common drivers of lethal aggression are predation and territorial defense. In northeastern Poland, the yellow-necked mouse (Apodemus flavicollis) coexists with several rodent species, including the bank vole (Clethrionomys glareolus). Compared to voles, A. flavicollis is larger, physically stronger, more aggressive, and dominant in the social ecosystem. However, no visually documented instance of a lethal attack by this species has been reported up to date. Here, we present the first recorded case of a fatal attack by a yellow-necked mouse following an encounter with a bank vole. A near-lethal attack is also reported. Importantly, these attacks were not predatory, as no consumption occurred. The attacks appeared instead to be related to interspecies competition, i.e., to competitive interactions between two species that live in the same habitat and use the same type of resources. Notably, while the aggressiveness of yellow-necked mice towards bank voles was known, it was unknown that it could take such extreme forms. Since, in rodents, most competition-related agonistic interactions are aimed at distancing the competitor, the physical destruction of the competitor appears as a surprisingly extreme way of addressing the game of interspecies competition through definitive removal of the opponent. Our observations highlight the need for further research on interspecific aggression among small mammals. They also emphasize the importance of field-based methods, such as camera trapping and continuous video monitoring, which allow for direct observation of animal behavior in natural settings and can reveal rare or previously overlooked interactions.

Approaching conspecifics is a fundamental component of social interactions. Such decisions are typically investigated through the study of social relationships (e.g., kinship, dominance, affiliation) and their impact. In the framework, the role of social affordances, i.e., interaction opportunities arising from the immediate social environment, remains poorly understood. We investigated how context-dependent factors shape approach decisions in wild crested macaques (Macaca nigra) in North Sulawesi, Indonesia. From September 2024 to May 2025, we collected data on 27 adults (20 females, 7 males) using focal observations and an approach-specific protocol recording neighbours identities and behaviours at the initiation of close-proximity approaches (<1 body length). We first compared data collected during approaches with focal data, then conducted within-approach analyses contrasting approached partners and bystanders. We further tested the effects of behavioural state, subgroup size and composition, including infant presence. Bayesian models revealed that affiliative, neutral, and grooming behaviours were more frequent in approach contexts than in general observations. Within approaches, individuals avoided congeners already engaged in grooming and approached individuals in neutral or affiliative states (e.g., lip-smacking, embracing). Macaques globally approached more isolated individuals or dyads. When multiple subgroups were present (i.e., multiple choices), the likelihood of being approached increased with subgroup size and was higher in subgroups containing infants. These findings indicate that macaques use behavioural cues and local social density as affordance signals, adjusting approach decisions to maximize interaction opportunities and navigate complex social environments.

Zoological gardens strive to prioritize excellent animal care and adhere to the highest standards of animal welfare, ensuring that the conditions of animals` social and physical environment are as close as possible to those in the wild. This study investigates the effects of group split and relocation to the new enclosure on the behaviour of western lowland gorillas (Gorilla gorilla gorilla) (N = 7) living in Prague Zoo, Czech Republic. We conducted over 289 hours of behavioural observations focusing on daily activities, social interactions, and behaviours that serve as potential stress and welfare indicators. The group split led to establishment of two groups in two separate enclosures; the old enclosure consisted of a "bachelor group" (i.e. three males) and a new enclosure consisted of a mixed-sex group (i.e. three females and one juvenile male). The behavioural comparisons across different study periods were conducted using linear mixed models (LMMs). The changes led to an increase in time spent moving, feeding, being in social proximity, and higher rates of approaches among the gorillas, as well as to a decrease in rates of self-directed and "undesirable" behaviors. Our findings indicate that the gorillas effectively adapted to the changes, most likely by relying on social support, to navigate new conditions. This study contributes to our understanding of how socio-cognitively complex species cope with necessary alterations in captive animal care programs. Furthermore, these observations may inform strategies to enhance the welfare of zoo-housed animals and to improve their captive care.

Dog (Canis lupus familiaris) vocal communication occurs across a wide range of social contexts, including dog-human social play, a common and dynamic interaction in which diverse vocal signals are expressed, particularly in young dogs. However, most available open datasets of dog vocalizations focus primarily on barks, leaving other sound types underrepresented. Here we present a bioacoustic dataset of dog-human social play sessions recorded under standardized laboratory conditions, comprising 30 play sessions involving 17 young dogs (6-24 months old) of different breeds playing with familiar humans. Raw audio recordings are accompanied by two layers of annotations covering the dog sounds produced during sessions. Video recordings of the sessions are provided for contextual reference, along with metadata describing each dog and the experimental sessions. Additionally, permutation-based classification analysis showed that annotated sound categories exhibit above-chance and generalizable acoustic differences across individuals. The dataset may support research on dog vocal communication and expand the range of sounds documented during positive dog-human social interactions.

The Clouded Tiger Cat Leopardus pardinoides is a recently recognized Neotropical species for which ecological and natural history data are sparse. Knowledge of species distribution and elevational range are largely based upon camera trap studies, and its diet has not been examined. The objective of this study was to better define the geographical and elevational distribution of the subspecies L. pardinoides oncilla in Central America using genetically confirmed records. We also provide the first diet analysis for L. pardinoides. We conducted extensive surveys for scat samples by visual means and with a scent detection dog across mountain ranges in Panama and Costa Rica, confirmed species identity using Sanger sequencing of mitochondrial DNA, and analyzed diet composition via DNA metabarcoding. Altogether, we collected 195 confirmed L. pardinoides scats. Records were strongly associated with high elevations, with a median elevation of 2805 m, and a maximum of 3422 m. Our findings highlight core habitat for L. pardinoides in the Cordillera Talamanca mountains in the border region of Panama and Costa Rica, and we identified isolated populations from the Central Volcanic Cordillera of Costa Rica and the Central Cordillera of Panama. DNA metabarcoding detected 59 vertebrate taxa. Diet was dominated by small mammals (74% of samples), especially cricetid rodents and shrews. Birds were also commonly detected (35% of samples), while reptiles (13%) and amphibians (1.9%) were less frequent. Estimated median adult prey mass was low (25 g), indicating that the species specializes on small prey. Most vertebrate prey identified were species endemic to montane regions of Panama and Costa Rica. These findings provide valuable ecological and natural history data on Clouded Tiger Cats in Central America that will inform conservation and management of this recently recognized species.

Mother-daughter relationships are central to social organization in rhesus macaques, providing the foundation for kin familiarity, affiliative bias, and intergenerational rank inheritance. Understanding how these bonds are structured is therefore important for identifying the processes that contribute to matrilineal cohesion and for anticipating risks of social instability and matriline fragmentation in captive groups. We studied 141 dam-daughter dyads in large, captive groups of rhesus macaques, using generalized linear models to examine associations between demographic variables and affiliative behaviors, and a multivariate clustering approach to characterize how these behaviors combine into distinct relationship types. Our analyses identified eight distinct dam-daughter relationship types defined by combinations of three behavioral domains: grooming, spatial closeness, and agonistic support. Generalized linear models showed that demographic variables had domain-specific effects, with family size, age gaps, and dam age differentially associated with grooming, proximity, and support. These patterns were reflected in the clustering analysis, in which dyads differed not only in overall levels of behavior but also in how behaviors combined within dyads. For example, some dyads exhibited high grooming and support despite low spatial proximity, whereas others showed consistently high expression across all domains. Together, these results demonstrate that dam-daughter bonds are multidimensional and structured through the combination of multiple behavioral domains influenced by demographic and life-history factors. Because these dyads scaffold interactions among close kin, variation in dam-daughter bond structure may influence the development and stability of broader matrilineal relationships. From an applied perspective, considering multiple behavioral domains when assessing social relationships can help identify dyads that may warrant closer monitoring and inform management strategies to support matrilineal cohesion and reduce the risk of social instability in captive groups.

For many group living species, vocal signals are a vital form of communication needed to coordinate social behaviours. Investigating these processes first requires a detailed description of the species vocal repertoire. White-nosed coatis (Nasua narica), which forage and move together in forest habitats, are thought to rely on vocalisations to coordinate movements and maintain group cohesion. However, quantitative studies of their vocal repertoire are lacking. We examined the vocal repertoire of white-nosed coatis from wild populations in Panama and Arizona, USA, to gain a more comprehensive view of their calling behaviours. By combining traditional acoustic analyses with an unsupervised approach based on spectrogram structure, we characterised the diversity of calls in this species and described the temporal and structural features of their vocalisations. We identified 19 call types, with some of these calls emitted in multi-syllable call sequences or in fast succession. We found variability in call rates among group members, which may be driven by differences in social status within the group. In addition, our results indicate that white-nosed coatis likely have individually-recognisable vocalisations. This study provides a foundational description of the white-nosed coati vocal repertoire, laying the groundwork for future research on vocal communication in this species.

Vocal communication depends not only on the acoustic features and rate of vocalizations, but also on their timing relative to the vocalizations of others. In this study, we examined whether ultrasonic vocalizations (USVs) produced by adult female Mongolian gerbils during freely moving social interactions are temporally related to the vocal timing of a partner. Using USVCAM, we assigned each USV to an individual caller and evaluated whether brief USVs exhibit turn-taking-like temporal organization by analyzing response latency after partner calls, call overlap rate, and circular-shift surrogate data. The offset-to-onset gaps of alternating vocalizations were concentrated within a short time range, with a median of 130 ms. In addition, the call overlap rate was significantly lower in the observed data than in the circularly shifted surrogate data, indicating that USVs occurred with short latencies after partner calls while being less likely to overlap with ongoing partner calls. Furthermore, USV counts were reduced in unoperated Ctrl animals that interacted with devocalized Mute animals. These findings suggest that gerbil USVs may be coordinated in relation to partner vocal timing and vocal input. This study provides a basis for understanding rodent USVs not merely as individual vocal outputs, but as temporally organized dyadic social signals.

Understanding how anthropogenic disturbance and vegetation structure influence bird abundance is important for biodiversity conservation in rapidly changing tropical landscapes. This study evaluated the effects of anthropogenic and vegetation-related variables on the abundance of the Red-cheeked Cordon-bleu (Uraeginthus bengalus) in human settlements and surrounding farmlands in Laminga Village, Jos-East Local Government Area, Plateau State, Nigeria. Bird surveys were conducted using line transects and quadrat-based vegetation assessments during November 2024. Poisson Generalized Linear Models (GLMs) were used to examine the influence of anthropogenic and vegetation predictors on abundance. Among anthropogenic variables, building density significantly reduced abundance ({beta} = -0.141, SE = 0.060, z = -2.333, p = 0.020), whereas human presence ({beta} = -0.073, p = 0.141) and noise level ({beta} = 0.009, p = 0.592) did not significantly influence abundance. Average grass height showed a marginal positive relationship with abundance ({beta} = 2.008, SE = 1.051, z = 1.910, p = 0.056), while hedgerow presence, hedgerow height, grass cover, and bare ground cover were not significant predictors. The vegetation model produced the lowest residual deviance (91.19) and AIC value (297.66), indicating comparatively stronger explanatory performance. The results suggest that structural habitat characteristics and building density may play more important roles in shaping Red-cheeked Cordon-bleu abundance than human activity or noise levels alone. These findings provide insight into species responses to environmental disturbance in human-modified savanna ecosystems.

Brydes whales (Balaenoptera edeni edeni) are nationally protected in China, and the waters around Weizhou Island in the Beibu Gulf support one of the countrys few regularly observed coastal groups. However, acoustic data for this population remain limited, and the potential effects of local vessel noise are poorly described. We conducted 16 vessel-based surveys around Weizhou Island and adjacent waters in January 2024 using a low-disturbance sailboat platform and passive acoustic recorders, with concurrent visual observations where possible. We identified 734 low-frequency signals classified as putative Brydes whale vocalizations and quantified their temporal and spectral parameters. Call duration was significantly negatively correlated with maximum frequency and center frequency, but not with minimum frequency or bandwidth. Comparisons with published records indicate that the recorded signals are most similar to vocalizations previously reported from juvenile Brydes whales or mother-calf pairs, although individual source attribution could not be confirmed. Speedboat passage significantly increased root-mean-square sound pressure levels, and the dominant noise band overlapped the frequency range of the recorded Brydes whale signals, indicating potential for acoustic masking. These results expand the bioacoustic baseline for Brydes whales in Chinese coastal waters and provide evidence relevant to the management of vessel activity and whale-watching tourism around Weizhou Island.

As wolves (Canis lupus) recolonize diverse environments, the extent to which ecological and individual factors shape their spatial behaviour remains insufficiently understood. Italy, where wolf recovery began earlier than elsewhere in Europe and spans highly heterogeneous landscapes, offers an opportunity to explore patterns relevant to future European scenarios. We analysed variation in home range size and movement patterns using GPS data from 19 wolves monitored between 2019 and 2025 across five study areas representing a broad environmental gradient. Continuous-time movement modelling was used to estimate home range size and distance travelled. Home range size varied markedly among areas, with annual averages for pack-member wolves ranging from 35 to 185 km2. The smallest ranges occurred where wolves had been established for longer and/or where prey availability was higher, suggesting that both intraspecific competition and resource abundance influence space use. Wolves expanded their ranges in winter and in more urbanized landscapes. Pack members consistently had smaller home ranges than floaters, and among floaters, females showed slightly larger ranges than males. Distance travelled also differed significantly across areas (local averages 25-53 km/day) but showed no significant association with individual traits. Three dispersal events were documented, two of which resulted in successful pair formation and subsequent reproduction. Overall, our results provide a broad overview of wolf spatial behaviour in contemporary Italian landscapes and offer insights relevant to the management of wolf populations across Europe.

Common terns (Sterna hirundo) are regionally threatened migratory seabirds that form large breeding colonies during the North American summer months. They are highly vocal and serve as important bioindicators of aquatic ecosystems. Historically, acoustic studies on colonial seabirds have proven difficult due to the dense aggregations of individuals and high rate of call overlap. However, as passive acoustic monitoring (PAM) becomes increasingly common for studying seabird colonies, quantitative descriptions of species vocalizations are needed to accurately interpret behavioral information from colony soundscapes and support automated analysis of large acoustic datasets. This study aims to quantify the vocal repertoire of adult common terns. We deployed AudioMoths to collect acoustic data at a tern colony on Seavey Island, New Hampshire, USA from across the breeding season. Using RavenPro, unique call types were identified through visual and aural inspection of the acoustic data in the spectrogram. For each call, we then extracted measurements of peak frequency (Hz), bandwidth 90% (Hz), syllable duration 90% (s), and total bout duration (s) to quantify the characteristics of each call type. Statistical analyses for acoustic parameters by call type were performed using Kruskal-Wallis tests, followed by post-hoc Dunn tests. Our results demonstrate that each call type is significantly different from another by at least one parameter, with the exception of the kek and kip/tjuk calls. These findings present the first quantitative analysis of common tern vocalizations for North America. By defining temporal and spectral characteristics for multiple call types, this work helps translate colony soundscape into biologically meaningful information about tern behavior and colony dynamics. These descriptions also provide key parameters for developing automated tools to detect and classify vocalizations in dense, noisy colonies. Integrating quantified vocal characteristics with PAM offers a promising approach for monitoring colony activity and behavior while minimizing disturbance relative to traditional methods.

Non-human primates are crucial animal models in biomedical research including neurosciences. Legal frameworks ensure that participation of monkeys in biomedical research follows an ethical assessment, driven by specific guidelines for ensuring animal well-being. Those guidelines promote continued refinement of practices to improve animal wellbeing, yet quantitative data to inform such refinements is limited. In this context, we investigated the impact of modifications to the housing that created either navigational complexity or visual access to neighbouring animals on the behaviour of rhesus macaques. Then we quantified the value of different enrichment programmes based on manipulable objects. We showed that making navigation within a modular housing more complex, or adding transparent separations between housing units is associated with significant reduction of severe aggression, stereotypical behaviours and a significant increase of appeasing behaviours. We also observed a positive effect of manipulable objects with an increase in the expression of appeasing behaviours and a decrease in abnormal or stereotypical behaviours. Finally, our study revealed significant inter-individual variability that was not explained by sex, age, social network size or social status. Overall, our results point to the importance and benefits of environmental changes and enrichment protocols, and of the critical need for personalised enrichment programmes.

The human larynx, compared to those of closely related primates, lies deeper in the throat and lacks vocal membranes and air sacs. These shifts are usually analyzed regarding their acoustic effects on vowel-like vocalizations, since the evolution of speech was long thought to require an expansion of vocal range driven by vocal tract modifications. However, vowels are just one type of phoneme, and speech is just one class of human utterance. To understand the evolutionary underpinnings of known shifts in human vocal morphology, a broader bioacoustic comparison is needed. Specifically, the range of sounds used in human speech must be compared to that employed in other human vocalizations and in the repertoires of extant close primate relatives. Here, we measure the acoustic-feature space occupied by human speech, non-linguistic, and musical vocalizations along with the calls of chimpanzees, bonobos, and chacma baboons. We use Mel-frequency cepstral coefficients to create an acoustic space depicting the spectro-temporal features of over 750,000 brief vocal segments sourced from published databases and other verified sources. Speech and song occupied significantly less volume in this acoustic space than human non-linguistic vocalizations. In addition, the acoustic-feature volumes of speech and song were not statistically distinct from those of non-human primates. These results suggest that speech was not enabled by an expansion of human vocal acoustic space. Anatomical shifts unique to humans may have led to an elaboration of non-linguistic utterances, but learned vocalizations use a surprisingly small fraction of this space. Our understanding of human vocal evolution will be further informed by additional systematic comparisons of the function and homology of non-speech vocalizations, along with the collection and incorporation of more complete non-human primate vocal datasets, especially from Gorilla and Orangutan.

Animal vocalisations can convey information about external events, but whether this goes beyond reflecting the emotional state elicited by these events is debated. To explore this, we studied the acoustic structure of common marmoset (Callithrix jacchus) phee (long-distance contact) and ek (alert/mobbing) calls produced in five treatments varying in the emotional valence and arousal they elicit (internal state), as well as food and social context (external events). We measured changes in arousal via nasal temperature and analysed both basic acoustic parameters and Mel-frequency cepstral coefficients (MFCCs) of the calls. Support Vector Machines combined with Linear Mixed effect models revealed that phee calls encode both external events and internal states, while eks reflected predominantly arousal. Notably, an acoustic signature related to food context was present in phees both when provided (positive valence) and teased with highly preferred food items (negative valence), and even when food was not physically present (food call playback treatment). This suggests marmoset long-distant phee calls encode external information beyond emotional arousal and valence, and independently of the presence of an immediately triggering stimulus.

Rhesus macaques (Macaca mulatta) are widely used as non-human primate models for biomedical research. When housed in captivity, it is essential to provide an environment that supports their natural behaviours; otherwise, they risk developing mood disorders, stereotypies, and other behavioural issues that may lead to physical harm. The objective of this preliminary study was to monitor the behaviour of three aged rhesus macaques ([&ge;] 20 y.o.), relocated from a laboratory to a Rescue Centre for Exotic Animals (Italy), and to assess the impact of novel food enrichments. Behavioural data were collected over 18 weeks, beginning at their arrival, using continuous focal sampling from video recordings. Simultaneously, faecal samples were gathered for cortisol analysis. The study was divided into three phases: a control phase without enrichments, a feeding enrichment phase (divided into two periods), and a final control phase without enrichments. Each phase comprised 900 minutes of observations for each subject. Data were analysed using generalized linear mixed models. Results showed an increase in locomotion during the enrichment and final phase compared to the initial phase. Additionally, a reduction in scratching and body-shaking behaviours was observed in the final phase compared to the initial phase. These findings suggest that implementing an enrichment program can enhance the welfare of aged non-human primates and can be considered a valuable tool in the rehabilitation of non-human primates previously housed in laboratories. GRAPHICAL ABSTRACT O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=113 SRC="FIGDIR/small/719840v1_ufig1.gif" ALT="Figure 1"> View larger version (50K): org.highwire.dtl.DTLVardef@152a3a1org.highwire.dtl.DTLVardef@74b53forg.highwire.dtl.DTLVardef@275b21org.highwire.dtl.DTLVardef@1d004d8_HPS_FORMAT_FIGEXP M_FIG C_FIG RESEARCH HIGHLIGHTSO_LIEnvironmental enrichment positively affected activity and stress indicators in aged ex-laboratory rhesus macaques. C_LIO_LILocomotion rates increased while scratching, body-shaking, and cortisol levels decreased. C_LIO_LIEnrichment enhance welfare during rehabilitation, even in older individuals. C_LI

AimHome range size is a fundamental aspect of animal spatial ecology, and understanding the factors that shape it is important for conservation purposes. Several hypotheses, based on energy needs or competition, assume that home range size negatively correlates with population density. However, this pattern has been little investigated on a global scale, and it remains unclear whether it would stand at both intra- and interspecific levels. To fill this gap, we conducted a global exploration of this relationship at the level of an animal family. Location: Global. Time period: Contemporary. Major taxa studied: Wild Felidae. MethodsIndividual home range size records (n = 1022) and population density estimates (n = 1061) were retrieved from the literature for 23 felid species across the world. We first investigated the interspecific relationship by modelling the median home range size of a species as a function of its median population density. To study the intraspecific relationship, we spatially merged data points based on their spatial or temporal proximity. We then applied a mixed-effects linear model using species as a random factor. ResultsWe found that home range size was negatively associated with population density, at both interspecific (-1.323 {+/-} 0.180, p < 0.001) and intraspecific levels (-0.569 {+/-} 0.201 to - 0.537 {+/-} 0.201 depending on the merging approach, p < 0.01). Landscape features were also predictors of home range size, without confounding the effect of population density. Main conclusionsSeveral processes likely govern the relationship between home range size and population density: differences in body mass between species may drive the interspecific relationship, whereas the intraspecific pattern is probably explained by conspecific competition. Although more research is needed to quantify their relative contribution, our study highlights a worldwide ecological pattern that exists at multiple biological levels in the wild.

Risk-taking behaviour permeates daily decision-making across many taxa and has significant impact on fitness outcomes. Previous research finds that individuals display consistent tendencies in their propensity to take risks, often labelled as a "Boldness-Shyness" personality trait. In this experimental study, we investigated repeatability in risk-taking behaviour in 241 wild vervet monkeys by measuring their responses to capture threat (via human-initiated darting procedure) and predation threat (via two predator models). We further examined the influence of socio-demographic, group identity and hormonal factors on risk-taking behaviour. We found that multiple behavioural responses, including the likelihood of approaching a food source under threat, were consistent across contexts. Risk-taking behaviour was influenced by age, sex, dominance rank, and hormonal profiles: juveniles, males, and higher-ranking individuals were more likely to approach food under (perceived) predation risk. Additionally, we observed significant among-group differences, suggesting that individuals within the same social group exhibit similar risk sensitivities, potentially due to social facilitation or shared environmental effects. The findings contribute to our understanding of animal personality and the ecological and social drivers underlying variation in risk-related behaviours.

The diversity of the brown frog genus Rana may be underestimated as the high similarity of morphological characters. A new species belonging to the genus Rana is delineated based on eight specimens obtained from the Tianma National Nature Reserve, Jinzhai County, Luan City, Anhui Province, China. The phylogenetic analysis based on three mitochondrial genes (12S, ND2, and Cyt b) and one nuclear gene (BDNF) showed that the new species formed an independent clade closely related to R. culainensis and received strong support. In addition, morphological differentiation confirmed the phylogenetic results, and both supported the validity of a new species (Rana tianmaensis sp. nov.) in the R. japonica species group. The discovery of this new species enhances peoples understanding of the biodiversity of Rana and can provide important foundational data for scientific decision-making on protected area construction, ecological conservation, and species diversity. With the inclusion of newly described species in this study, the distribution of Rana genus in China now includes 31 recognized species.