Svetlana S. Gogoleva

TO BARK OR NOT TO BARK: VOCALIZATIONS BY RED FOXES SELECTED FOR TAMENESS OR AGGRESSIVENESS TOWARD HUMANS

ABSTRACT In this study we classify call structures and compare vocalizations toward humans by captive red foxes Vulpes vulpes, artificially selected for behaviour: 25 domesticated, or “Tame” animals, selected for tameness toward people, 25 “Aggressive” animals, selected for aggression toward people, and 25 “Unselected” control foxes, representing the “wild” model of vocal behaviour. In total, 12,964 calls were classified visually from spectrograms into five voiced (tonal) (whine, moo, cackle, growl and bark), and three unvoiced, or noisy (pant, snort and cough) call types. The classification results were verified with discriminant function analysis (DFA) and randomization. We found that the Aggressive and Unselected foxes produced the same call type sets toward humans, whereas the Tame foxes used distinctive vocalizations toward humans. The Tame and Aggressive foxes had significantly higher percentages of time spent vocalizing than the Unselected, in support of Cohen & Fox (1976) hypothesis that domestication relaxes the selection pressure for silence, still acting in wild canids. Unlike in dogs, the “domesticated” Tame foxes did not show hypertrophied barking toward humans, using instead the cackle and pant. We conclude that the use of a certain call type for communication between humans and canids is species-specific, and not is the direct effect of domestication per se.

Sign and strength of emotional arousal: vocal correlates of positive and negative attitudes to humans in silver foxes ( Vulpes vulpes )

Summary The hypothesis of similarity in trends of acoustic characteristics regardless of the sign of emotional arousal, positive or negative, has been advanced based on human vocalizations. For non-human mammals, testing is complicated because the same stimulus cannot evoke opposite (positive and negative) internal states, to trigger the respective vocalizations. To resolve this concern, we designed an experimental procedure using Tame and Aggressive strains of silver foxes, with genetically predetermined positive or negative emotional responses to humans respectively. We analyzed features of vocalizations produced by callers at different fox‐human distances, assuming changes in vocal responses reflect the shifts of human-related positive arousal in Tame foxes and human-related negative arousal in Aggressive foxes. Both strains showed similar trends for changes in calling rate and proportion of time spent vocalizing toward higher levels in response to greater emotional arousal, positive in Tame foxes and negative in Aggressive foxes. At the same time, strains showed distinctive trends for the proportions of different call types and maximum amplitude frequency. We infer that the variables with similar trends reflect the strength of emotional arousal, regardless of triggering internal states, whereas variables with distinctive trends are specifically related to the sign of emotion in silver fox.

Kind granddaughters of angry grandmothers: The effect of domestication on vocalization in cross-bred silver foxes

The genetic basis of the effects of domestication has previously been examined in relation to morphological, physiological and behavioural traits, but not for vocalizations. According to Belyaev [Belyaev, D.K., 1979. Destabilizing selection as a factor in domestication. J. Hered. 70, 301-308], directional selection for tame behaviour toward humans resulted in domestication. This hypothesis has been confirmed experimentally on the farm-bred silver fox Vulpes vulpes population that has undergone 45 years of artificial selection for tameness and 35 years of selection for aggressiveness. These foxes, with their precisely known attitudes toward people, provide a means of examining vocal indicators of tameness and aggressiveness to establish the genetic basis for vocal production in canids. We examined vocalizations toward people in foxes selected for tameness and aggressiveness compared to those of three kinds of crosses: Hybrids (Tame x Aggressive), A-Backcrosses (Aggressive x Hybrid) and T-Backcrosses (Tame x Hybrid). We report the effects of selection for tameness on usage and structure of different vocalizations and suggest that vocal indicators for tameness and aggressiveness toward people are discrete phenotypic traits in silver foxes.

The gradual vocal responses to human-provoked discomfort in farmed silver foxes

Vocal indicators of welfare have proven their use for many farmed and zoo animals and may be applied to farmed silver foxes as these animals display high vocal activity toward humans. Farmed silver foxes were selected mainly for fur, size, and litter sizes, but not for attitudes to people, so they are fearful of humans and have short-term welfare problems in their proximity. With a human approach test, we designed here the steady increase and decrease of fox–human distance and registered vocal responses of 25 farmed silver foxes. We analyzed the features of vocalizations produced by the foxes at different fox–human distances, assuming that changes in vocal responses reflect the degrees of human-related discomfort. For revealing the discomfort-related vocal traits in farmed silver foxes, we proposed and tested the algorithm of “joint calls,” equally applicable for analysis of all calls independently on their structure, either tonal or noisy. We discuss that the increase in proportion of time spent vocalizing and the shift of call energy toward higher frequencies may be integral vocal characteristics of short-term welfare problems in farmed silver foxes and probably in other captive mammals.

Two new species of Kalophrynus Tschudi, 1838 (Anura: Microhylidae) from the Annamite mountains in southern Vietnam.

We describe two new mountain-dwelling microhylid species of the genus Kalophrynus from the southern part of the Annamite Mountains in Vietnam. The two new species differ from all known congeners in morphological characters and mtDNA; phylogenetically, they form a sister clade to the large-bodied K. interlineatus (1009 bp, 16S rRNA gene, mtDNA). Both species share the following characteristics: snout pointed in dorsal and lateral views, slightly sloping in profile; tympanum distinct, smaller than eye in diameter; toe webbing moderate; outer metatarsal tubercle present; light dorsolateral line absent. Kalophrynus cryptophonus sp. nov. from Loc Bao, Lam Dong Province is a small-sized species distinguishing from its congeners by a combination of: SVL 27.9-30.4 mm in males, 23.4 mm in female; canthus rostralis indistinct; males with large sharp conical spines on the skin covering mandible margins and finely asperous nuptial pads on the dorsal surface of fingers I-III; dark ocelli in the inguinal region usually present, small, without a light border; anterior palatal dermal ridge short, restricted to medial part of palate. Kalophrynus honbaensis sp. nov. from Hon Ba, Khanh Hoa Province is a medium-sized Kalophrynus, distinguishing from its congeners by a combination of: SVL 26.7-36.8 mm in males; canthus rostralis distinct; males without distinguishable spines on the mandible margins nor the nuptial pads; dark ocelli in the inguinal region present, large, without a distinct light border, anterior palatal dermal ridge developed, parallel to posterior one. Kalophrynus cryptophonus sp. nov. reproduces in hollow bamboo stems; we describe larval morphology and bioacoustics of this species in relation to phytotelm breeding. A review of the distribution of the genus Kalophrynus in Indochina is provided.

Explosive vocal activity for attracting human attention is related to domestication in silver fox.

Domestication affects behavioral and vocal responses, involved in communication with humans; in particular, those that attract human attention. In this study, we found that silver foxes of Tame strain, experimentally domesticated for a few tenses of generation, displayed bursts of vocal activity during the first minute after appearance of an unfamiliar human, that faded quickly during the remaining time of the test, when the experimenter stayed passively before the cage. Distinctively, foxes of Aggressive strain, artificially selected for tenses of generation for aggressive behavior toward humans, and the control group of Unselected for behavior silver foxes kept steady levels of vocal activity for the duration of the tests. We found also that Aggressive foxes vocalized for a larger proportion of time than Unselected foxes for all 5 min of the test. We discuss the obtained data in relation to proposal effects of domestication on mechanisms directed to involving people into human-animal interactions and structural similarity between human laughter and vocalization of Tame foxes.

Vocalization toward conspecifics in silver foxes (Vulpes vulpes) selected for tame or aggressive behavior toward humans

We examined the production of different vocalizations in three strains of silver fox (unselected, aggressive, and tame) attending three kinds of behavior (aggressive, affiliative, and neutral) in response to their same-strain conspecifics. This is a follow-up to previous experiments which demonstrated that in the presence of humans, tame foxes produced cackles and pants but never coughed or snorted, whilst aggressive foxes produced coughs and snorts but never cackled or panted. Thus, cackle/pant and cough/snort were indicative of the tame and aggressive fox strains respectively toward humans. Wild-type unselected foxes produced cough and snort toward humans similarly to aggressive foxes. Here, we found that vocal responses to conspecifics were similar in tame, aggressive and unselected fox strains. Both cackle/pant and cough/snort occurred in foxes of all strains. The difference in the use of cackle/pant and cough/snort among these strains toward humans and toward conspecifics suggest that silver foxes do not perceive humans as their conspecifics. We speculate that these vocalizations are produced in response to a triggering internal state, affiliative or aggressive, that is suppressed by default in these fox strains toward humans as a result of their strict selection for tame or aggressive behavior, whilst still remaining flexible toward conspecifics.

Effects of selection for behavior, human approach mode and sex on vocalization in silver fox

This study presents a first direct comparison of vocal type, call rate and time spent vocalizing among Unselected, Tame and Aggressive strains of silver fox (Vulpes vulpes) in three modes of human approach (Provoking, Approach–Retreat, and Static). Also, it provides a first comparison of male and female vocal output in the Provoking test. Vocal types were found strain-specific irrespective of the fox sex or the test. Males had higher call rates and spent shorter times vocalizing than females. These results support the evidence of genetic-based emotional states, triggering vocal behavior in silver fox strains, and suggest sex dimorphism in vocal activity toward humans.

Molecular, morphological and acoustic assessment of the genus Ophryophryne (Anura, Megophryidae) from Langbian Plateau, southern Vietnam, with description of a new species

Abstract Asian Mountain Toads (Ophryophryne) are a poorly known genus of mostly small-sized anurans from southeastern China and Indochina. To shed light on the systematics within this group, the most comprehensive mitochondrial DNA phylogeny for the genus to date is presented, and the taxonomy and biogeography of this group is discussed. Complimented with extensive morphological data (including associated statistical analyses), molecular data indicates that the Langbian Plateau, in the southern Annamite Mountains, Vietnam, is one of the diversity centres of this genus where three often sympatric species of Ophryophryne are found, O. gerti, O. synoria and an undescribed species. To help resolve outstanding taxonomic confusion evident in literature (reviewed herein), an expanded redescription of O. gerti is provided based on the examination of type material, and the distributions of both O. gerti and O. synoria are considerably revised based on new locality records. We provide the first descriptions of male mating calls for all three species, permitting a detailed bioacoustics comparison of the species. We describe the new species from highlands of the northern and eastern Langbian Plateau, and distinguish it from its congeners by a combination of morphological, molecular and acoustic characters. The new species represents one of the smallest known members of the genus Ophryophryne. At present, the new species is known from montane evergreen forest between 700–2200 m a.s.l. We suggest the species should be considered Data Deficient following IUCN’s Red List categories.

越南中部发现棱皮树蛙属一新种（两栖纲：树蛙科）

A new species of small tree frog from a primary montane tropical forest of central Vietnam, Tay Nguyen Plateau, is described based on morphological, molecular, and acoustic evidence. The Golden Bug-Eyed Frog, Theloderma auratum sp. nov., is distinguishable from its congeners and other small rhacophorid species based on a combination of the following morphological attributes: (1) bony ridges on head absent; (2) smooth skin completely lacking calcified warts or asperities; (3) pointed elongated tapering snout; (4) vocal opening in males absent; (5) vomerine teeth absent; (6) males of small body size (SVL 21.8–26.4 mm); (7) head longer than wide; ED/SVL ratio 13%–15%; ESL/SVL ratio 16%–20%; (8) small tympanum (TD/EL ratio 50%–60%) with few tiny tubercles; (9) supratympanic fold absent; (10) ventral surfaces completely smooth; (11) webbing between fingers absent; (12) outer and inner metacarpal tubercles present, supernumerary metacarpal tubercle single, medial, oval in shape; (13) toes half-webbed: I 2–2¼ II 1½–2¾ III 2–3¼ IV 3–1½ V; (14) inner metatarsal tubercle present, oval; outer metatarsal tubercle absent; (15) iris bicolored; (16) dorsal surfaces golden-yellow with sparse golden-orange speckling or reticulations and few small dark-brown spots; (17) lateral sides of head and body with wide dark reddish-brown to black lateral stripes, clearly separated from lighter dorsal coloration by straight contrasting edge; (18) ventral surfaces of body, throat, and chest greyish-blue with indistinct brown confluent blotches; (19) upper eyelids with few (3–5) very small flat reddish superciliary tubercles; (20) limbs dorsally reddish-brown, ventrally brown with small bluish-white speckles. The new species is also distinct from all congeners in 12S rRNA to 16S rRNA mitochondrial DNA fragment sequences (uncorrected genetic distance P>8.9%). Advertisement call and tadpole morphology of the new species are described. Our molecular data showed Theloderma auratum sp. nov. to be a sister species of Th. palliatum from Langbian Plateau in southern Vietnam.