原題：NOTES AND OBSERVATIONS ON COURTSHIP AND MATING IN Tityus (Atreus) magnimanus POCOCK, 1897 (Scorpiones: Buthidae)?

作者：Ross LK

本文詳細(xì)記錄了碩螯戾蝎交配前后及過程中的各個細(xì)節(jié)，對于了解碩螯戾蝎交配中的不同動作含義有重要的參考意義！

ABSTRACT: Courtship and mating behaviors of the scorpion Tityus (Atreus) magnimanus are herein described, consisting of various components that pertain to four distinct behavioral stages. The courtship and mating rituals of Tityus (Atreus) magnimanus are similar to those of other scorpions. Behavioral components are presented in an ethogram to demonstrate their occurrence during mating sequences. The current report is presented as observational data that were acquired during life history studies of this species.

摘要：本文描述了碩螯戾蝎的求偶和交配行為，包括了處于四個不同行為階段的各種組成部分。碩螯戾蝎的求偶和交配儀式與其他蝎子相似。行為成分呈現(xiàn)在一個圖譜中，以展現(xiàn)它們在交配序列中的出現(xiàn)。目前的報告是在該物種的生命歷程研究中獲得的觀測數(shù)據(jù)。

INTRODUCTION

Courtship and mating among scorpion species, on which such data are available, comprise a series of complex, ritualized and characteristic behaviors that involve a courtship period of stereotyped acts leading up to sperm transfer from male to female via an external stationary spermatophore (1-5). Courtship in scorpions is usually divided into four distinct behavioral stages (different terminologies have been applied to individual stages by various authors): initiation, promenade á deux, sperm transfer and separation (3, 4, 6).

簡介

現(xiàn)有數(shù)據(jù)表明，蝎子的求偶和交配包括一系列復(fù)雜的、儀式化的獨特行為，這些行為包括求偶期的千篇一律的行為，這些行為導(dǎo)致精子通過外部固定的精莢從雄性轉(zhuǎn)移到雌性。蝎子的求偶通常分為四個不同的行為階段（不同作者對各個階段使用了不同的術(shù)語）：起始、漫步（注：該詞經(jīng)查為葡萄牙語“雙人漫步”，即公拉母游蕩的階段）、精子轉(zhuǎn)移和分離。

In the majority of scorpion species, courtship is initiated by males and typically involves a male approaching a female and grasping her chelae within his own (in some instances, a chelicerae-to-chelicerae grasp may be initiated by males and used simultaneously with the chelal grasp to control female movements). After the chelal grasp, the male initiates and guides the female in a series of coordinated movements termed promenade á deux. This courtship “dance” may last from five minutes to several days, with most courtship and mating sequences lasting from 30 to 60 minutes (3).

對大多數(shù)種類的蝎子而言是由雄性發(fā)起求愛行為，通常包括雄性接近雌性并抓住她的鉗子（在某些情況下，鉗子對鉗子的抓握可能是由雄性發(fā)起的，同時“鉗握”（注：應(yīng)為須肢緊握，為便于理解可譯為鉗握）用于控制雌性的動作）。在鉗握之后，雄性發(fā)起并引導(dǎo)雌性進(jìn)行一系列被稱為promenade á deux（雙人漫步）的協(xié)調(diào)動作。這種求偶“舞蹈”可能持續(xù)5分鐘到幾天，（注：一直以為持續(xù)幾天的情況是交配失敗了于是強制把公蝎拿出來打斷交配……圖樣圖森破）大多數(shù)求偶和交配過程持續(xù)30到60分鐘。

During the promenade, the male seeks out a suitable surface or structure upon which to deposit his spermatophore. Once the spermatophore has been extruded and attached to a suitable surface, the male then directs the female movements until her genital opening is positioned above the spermatophore. A brief struggle ensues and the female rocking motion assists in engaging the spermatophore with the genital operculum, and the sperm is released into this opening (7). Once sperm transfer is complete, the male usually initiates separation of the participants (3).

在“雙人漫步”的過程中，雄性會尋找一個合適的表面或結(jié)構(gòu)來放置精莢。一旦精莢被擠出并附著在合適的表面上，雄性就會引導(dǎo)雌性的運動，直到她的生殖器開口位于精子的上方。隨后是短暫的掙扎，雌性的搖擺動作幫助精莢與生殖器蓋接觸，精子被釋放到這個開口。一旦精子轉(zhuǎn)移完成，雄性通常會和雌性分開。

Courtship and mating behaviors have been described in approximately thirty scorpion species (3, 4), representing seven of the 18 extant families (8). Based on data reviewed by Polis and Sissom (3), both rituals have been described in more species (n = 12) from the family Buthidae than for any other scorpion family. Tityus C. L. Koch, 1836 is the largest genus within this family, represented by 181 species (9, 10). Tityus and the genus Centruroides Marx, 1890 (Buthidae), represent the only scorpion genera recognized as being medically significant in the Nearctic and Neotropical regions (11-13).

大約有30種蝎子被描述了求偶和交配行為，它們代表了現(xiàn)存18個科中的7個科。根據(jù)Polis和Sissom的數(shù)據(jù)，這兩種儀式在Buthidae科（殺牛蝎科/鉗蝎科）的物種中被描述的比在任何其他蝎子科中都多(n = 12)。戾蝎屬是該科最大的屬，共有181種（注：截止目前至少已有240種）。戾蝎屬和似刺尾蝎屬被認(rèn)為是在新北界和新熱帶界僅有的具有醫(yī)學(xué)意義的蝎子屬。

Despite the large number of species among Tityus, its abundance in many urbanized and rural human occupied areas, and the medical significance of many of its component species, courtship and mating in members of this genus are few in number (1, 14, 15).

盡管戾蝎屬的物種數(shù)量眾多，在許多城市化和農(nóng)村人類居住的地區(qū)都很豐富，而且其許多組成物種具有醫(yī)學(xué)意義，但該屬成員的求愛和交配數(shù)量很少。

The following contribution is based on laboratory observations of 12 mating sequences in the Venezuelan endemic scorpion, Tityus (Atreus) magnimanus Pocock, 1897 (Buthidae), conducted during February and March 2008. Tityus (Atreus) magnimanus inhabits tropical altitudinal forests in the states of Falcón and Lara in north-central Venezuela, and is considered to be of medical importance, with all severe and near-fatal envenomations in Falcón state attributed to this species. The present work is part of a continuing study on the life history of Tityus (Atreus) magnimanus and represents the first report on this medically significant species.

以下投稿基于2008年2-3月期間對委內(nèi)瑞拉特有的碩螯戾蝎的12個交配流程的實驗室觀察。碩螯戾蝎生活在委內(nèi)瑞拉中北部Falcón州和Lara州的熱帶高緯度森林中，被認(rèn)為具有重要的醫(yī)學(xué)意義，Falcón州所有嚴(yán)重和近乎致命的中毒都?xì)w因于該物種。目前的工作是對碩螯戾蝎生命周期的持續(xù)研究的一部分，是對這一醫(yī)學(xué)上重要物種的首次報道。

MATERIALS AND METHODS

In order to acquire female specimens of known age for life history studies, specimens of Tityus (Atreus) magnimanus were mated in the laboratory and observational data were recorded, where specimens of Tityus (Atreus) magnimanus (n = 84) were reared to maturity. Upon attaining sexual maturity, unrelated adult males (n = 12) and laboratory reared females (n = 12) were randomly selected and transferred to individual containers.

材料與方法

為了獲得已知年齡的雌性標(biāo)本用于生命周期研究，在實驗室對碩螯戾蝎樣本進(jìn)行了交配并記錄了觀察數(shù)據(jù)，飼養(yǎng)了84只碩螯戾蝎樣本至成熟。在性成熟后，隨機選擇不相關(guān)的成年雄性(n = 12)和實驗室飼養(yǎng)的雌性(n = 12)，并將其轉(zhuǎn)移到單獨的容器中。

Mature females were housed in circular 9 x 23 cm clear plastic containers, provided with a 50 mm layer of moist potting soil, a piece of cork or shelter and several flat rocks. As the containers of females would also serve as mating arenas, the addition of flat rocks provided a surface structure for spermatophore deposition by males.

性成熟的雌性被安置在9x23厘米的圓形透明塑料容器中，提供50毫米厚的潮濕盆栽土層、一塊軟木或其他遮蔽物、和幾塊平坦的巖石。由于雌性的容器也可以作為交配場所，所以增加的平坦巖石為雄性的精莢粘附提供了一個表面。

Adult males were kept in individual 15 x 15 x 15 cm clear plastic containers, with a 50 mm layer of moist potting soil and a single piece of cork for shelter. All specimens were maintained at temperatures from 26.6 to 29.0°C during the day and 23.8 to 25°C at night. The substrate was kept moist while ambient humidity levels were maintained in the range of 70 to 80%. Prey consisted of adult house crickets (Acheta domesticus L.) and cockroaches (Blatta lateralis Walker) with the type of prey alternated between ad libitum feedings. All observations were conducted under a low-intensity 60 watt red incandescent bulb in a parabolic reflector, suspended 50 cm above breeding arenas.

成年雄性被單獨飼養(yǎng)在15x15x15厘米的透明塑料容器中，容器中有50毫米的濕潤盆栽土壤和一塊軟木作為遮蔽物。所有樣本白天保持在26.6-29.0℃，夜間保持在23.8-25℃。環(huán)境濕度保持在70-80%的范圍內(nèi)，基底保持濕潤。食物以成體蟋蟀和櫻桃蟑螂為主，兩種食物隨意交替飼喂（注：ad libitum德語：隨意地）。所有的觀察都是在一個低強度的60瓦紅色白熾燈泡的拋物反射面下進(jìn)行的，該反射器懸掛在繁殖場所上方50厘米處。（注：蝎子對紅光不敏感，相關(guān)的資料正在學(xué)習(xí)如GREGORY R. C.等人關(guān)于不同波長光線對蝎子的活動影響（doi:10.1016/j.anbehav.2007.12.022）。也有Arachnoboard論壇上的網(wǎng)友表示蝎子根本看不見紅光，但我暫時沒找到相關(guān)文獻(xiàn)。）

Mature virgin females were provided a 24-hour acclimation period before mating trials were initiated. A randomly selected mature male was introduced into each of 12 breeding arenas and all observed courtship and mating behaviors were recorded.

在開始交配試驗之前，給予性成熟的處女雌性24小時的環(huán)境適應(yīng)期。在12個繁殖場所中，隨機選擇一只成年雄性，記錄所有觀察到的求偶和交配行為。（注：這只公可真是不虛蝎生?。?/p>

Once sperm transfer was completed, indicating termination of the mating sequence, males were removed from arenas and returned to individual containers.

一旦精子轉(zhuǎn)移完畢，這意味著交配過程結(jié)束了，公蝎從交配場所中被拿回單獨的容器里面。

RESULTS

In each of the 12 observed mating sequences, males immediately initiated courtship upon contact with females. As has been previously reported in scorpions (3), four distinct behavioral stages were observed: initiation, promenade á deux, sperm transfer, and separation. If specimens became separated during the promenade á deux, the behavioral sequence was immediately resumed by specimens without a return to a previous stage. The following ethogram demonstrates each behavioral component of courtship and the order in which they occurred during mating sequences (Figure 1).

結(jié)果

在12組交配進(jìn)程中，每組的公蝎見到母蝎后馬上就開始求愛，正如之前在參考文獻(xiàn)3中報道的一樣，觀察到四個不同的行為階段：起始、雙人漫步、精子轉(zhuǎn)移和分離。如果樣本在雙人漫步期間被分離，則樣本立即恢復(fù)繼續(xù)中斷的行為，而不會返回到前一階段。下表展示了求偶過程中每個行為組成部分以及它們在交配序列中發(fā)生的順序（表1）。

Stage I: Initiation

In all observed mating sequences, males initiated courtship after detecting the movements of females (n = 3; 25%) or during random encounters with stationary females (n = 9; 75%). When a male detected a moving female or encountered a stationary female, courtship was initiated by the male grasping and rapidly moving his chelae over the segments (including the chelae) of one (n = 8; 66.6%) or both (n= 2; 16.7%) pedipalps of a female.

第一階段:起始

在所有觀察到的交配序列中，雄性在發(fā)現(xiàn)雌性的運動后開始求愛(n=3；25%)或隨機遇到靜止的雌性(n=9；75%)。當(dāng)雄性發(fā)現(xiàn)一只移動的雌性或遇到一只靜止的雌性時，雄性就會抓住并迅速移動它其中一個鉗子(n=8；66.6%)或兩個（n=2；16.7%)。

Two males (16.7%) encountered immobile females and immediately grasped and moved their chelae over metasomal segments III to V of each female. After a brief period (less than 45 seconds) of metasomal or pedipalpal grasping behaviors, a male would grasp each chela of a female within his own and align his body with that of the female in a chelicerae-to-chelicerae position thus, making contact with the female and terminating the initiation stage of courtship.

2只雄性(16.7%)遇到靜止的雌性，立即抓住并將它們的鉗子移動到每一只雌性的第III至V節(jié)后體上方。在短暫的一段時間內(nèi)(少于45秒)的鉗住鉗子或后體的行為之后，雄性會抓住雌性的雙鉗，并將自己的身體與雌性以“鉗子對鉗子”的方式對齊，從而與雌性接觸，結(jié)束求愛的起始階段。

After initial contact by males, females typically withdrew their pedipalps against their own bodies and remained immobile until males initiated pulling movements. If a female resisted the attempts of a male to begin directed movements during initial contact, the male would judder for 10 to 25 seconds, then vigorously and repeatedly pull the female toward him until she began moving.

在與雄性進(jìn)行初次接觸后，雌性通常會將須肢縮回到自己的身體上并保持不動，直到雄性開始拉扯動作。如果雌性在初次接觸時拒絕雄性的引導(dǎo)動作，雄性會顫抖10到25秒，然后用力反復(fù)地將雌性拉向自己，直到雌性開始移動。

Polis and Sissom (3) reported that juddering by male scorpions functions either as a highly ritualized sexual or species-recognition behavior, as an instigator of mating behavior so that the female is stimulated to cooperate, or as a simple byproduct of intense sexual excitation. The only instances in which juddering was exhibited by males was in direct response to resistant females during the initiation and promenade á deux stages and appeared to be used by males to suppress unreceptive behaviors and stimulate females into mating.

Polis和Sissom報告說，雄蝎子的抖動，要么是一種高度儀儀化的性行為或是是一種物種識別行為——作為交配行為的煽動者，以此刺激雌性進(jìn)行協(xié)作交配，要么只是強烈性興奮的簡單表現(xiàn)。雄蟲表現(xiàn)出抖動的唯一情況是在起始期和“雙人漫步”期對雌性抵抗的直接反應(yīng)，似乎是雄蟲用來阻止雌性不接受的行為和刺激雌性交配。

In two pairings that involved receptive females that never resisted males during courtship movements, juddering was not observed in either male throughout all stages of courtship. These observations suggest that juddering behavior may be used by courting males to stimulate unreceptive females into mating.

在兩組求偶過程中從不抗拒雄性的雌性交配中，在求偶的所有階段都沒有觀察到雄性的抖動。這些觀察結(jié)果表明，抖動行為可能是求愛的雄性用來刺激不接受交配的雌性。

Alexander (1) reported metasomal grasping by courting males of Tityus (Atreus) trinitatis Pocock, 1897 and suggested that such behaviors by males may function as a means of identifying conspecific females through the sexually differentiated morphologies of metasomal segments. Repeated grasping of pedipalps and chelae of females by courting males may serve as a similar means of identifying conspecific females via differences in these morphological structures.

Alexander報道了求偶雄性碩螯戾蝎的“后體抓握”行為，并認(rèn)為雄性的這種行為可能是通過后體的性別分化形態(tài)來識別同種雌性的一種手段。雄性在向雌性求愛時反復(fù)抓住須肢和鉗子，可能是通過這些形態(tài)結(jié)構(gòu)的差異來識別同種雌性的一種類似手段。（注：通過網(wǎng)上各種信息的搜尋，國內(nèi)的蝎子飼養(yǎng)者普遍認(rèn)為蝎子的“控尾”——即文中的“后體抓握”行為是雄蝎讓雌蝎冷靜、保護自己的手段。個人比較贊同該觀點。因為仔細(xì)觀察，雌雄碰面甚至僅僅同處一缸感知到對方的存在之時，雙方有一個“發(fā)呆期”，此間雙方都在用櫛狀器拍打摩擦地面，甚至抖動身體來發(fā)送信號，此時應(yīng)該已經(jīng)完成了對對方物種的判斷，沒有必要再以生命為代價去識別后體。）

Sexually mature males and females of Tityus (Atreus) magnimanus exhibit differing morphologies of the pectines, chelae and metasomal segments. As has been reported by Louren?o (9), females of some species within the subgenus Atreus Gervais, 1843 possess enlarged basal middle lamellae that are not present in males. The chelal manus of males is moderately inflated and serves to differentiate mature males from immature males and females. Additionally, females are larger and more robust than males and the metasomal segments of sexually mature females are thickened and well developed in comparison to those of males.

性成熟的雌雄碩螯戾蝎的櫛狀器、鉗子和后體都表現(xiàn)出不同的形態(tài)。正如Louren?o所報道的那樣，1843年，一些Atreus亞屬（注：現(xiàn)為Scorpio，蝎屬）的雌性具有擴大的骨板中間片（注：即櫛狀器根部有“大齒”），而雄性沒有。雄性的鉗子有適度的膨脹，用以區(qū)分成熟雄性與不成熟雄性、雌性。此外，雌性比雄性更大、更健壯，性成熟雌性的后體比雄性更厚、更發(fā)達(dá)。

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Stage II: Promenade á Deux

The promenade á deux occurs in all scorpion species for which courtship and mating data are available (2, 3, 4). As in other scorpion species, the males of Tityus (Atreus) magnimanus direct and coordinate the movements of the pair during the promenade á deux. In Tityus (Atreus) magnimanus, the promenade begins when a male firmly grasps the chelae of a female within his own and begins to guide the female in mutually backward, sideways and rarely, forward movements. The male begins promenade movements by elevating its body slightly above the ground surface with the metasoma elevated and dorsomedially flexed, and moving backwards, effectively pulling the female along and directing her movements.

第二階段:雙人漫步

在所有有求偶、交配數(shù)據(jù)的蝎子物種中，雙人漫步都會發(fā)生。與其他蝎子物種一樣，雄性碩螯戾蝎在雙人漫步期間指導(dǎo)和協(xié)調(diào)配偶的行動。對碩螯戾蝎而言，當(dāng)一只雄性牢牢抓住一只雌性的螯，并開始引導(dǎo)雌性相互向后、向側(cè)面移動，很少向前移動時，漫步就開始了。雄性開始散步時，身體略高于地面，后體抬高并向背部彎曲，然后向后移動，有效地拉著雌性前進(jìn)，指導(dǎo)她的動作。

During backwards movements, the male will periodically flex and straighten the metasoma while simultaneously raising and lowering it in response to the willingness of the female to follow his movements.

在向后運動的過程中，雄性會周期性地彎曲和伸直后體，同時，為了讓雌性跟隨自己的運動，他們會根據(jù)雌性的意愿同步地抬起和放下后體。

If a female resists or stops moving, the male will move closer to her, elevate his body, and begin pulling the female in vigorous backward movements while simultaneously elevating and waving his metasoma from side-to-side. Females typically keep their bodies close to the substratum with the metasoma flexed and held laterally.

如果雌性抵抗或停止移動，雄性會靠近她，抬起身體，開始以有力的向后動作拉著雌性，同時抬起并左右擺動他的后體。雌性通常保持它們的身體靠近基底，而后體彎曲并保持在側(cè)面。

During this stage of courtship, the male guides the female across the substratum until a suitable surface is located for spermatophore deposition. During promenade movements, male pectines are directed downward and are moved back and forth over the substratum and all encountered surface structures. During promenade movements, the female may resist male attempts to direct coordinated movements of the pair by pulling and trying to move in a direction opposite that of the male or by ceasing movements, lowering her body to the substratum and becoming still.

在求愛的這一階段，雄性會引導(dǎo)雌性走過基底，直到找到一個適合存放精莢的表面。在雙人漫步中，雄性向下在基底和所有遇到的表面結(jié)構(gòu)上來回移動。在散步運動中，雌性可能會抵制雄性試圖引導(dǎo)其協(xié)調(diào)運動的嘗試，通過拉拽并試圖朝著與雄性相反的方向移動，或者停止運動，將身體降至基底并保持靜止。

Immobile females only resume moving after males exhibit juddering (6 to10 seconds) and vigorous pulling behaviors. In all observed pairings, the promenade á deux lasted from 4.5 to 9.0 minutes and was terminated when a suitable location for spermatophore deposition was selected by the male. A courting pair may pass over a structure several times from different directions until the male ceases their movements. Selection of a proper structure for spermatophore deposition finishes the promenade á deux.

靜止不動的雌性只有在雄性表現(xiàn)出抖動（6~10秒）和有力的拉扯行為后才能恢復(fù)活動。在所有觀察到的交配中，雙人漫步持續(xù)時間為4.5~9.0分鐘，當(dāng)雄性選擇合適的精莢放置位置時終止。一對求偶的蝎子可能會從不同的方向多次經(jīng)過一個基底，直到雄性停止它們的動作，為放置精莢選擇一個合適的結(jié)構(gòu)，雙人漫步才完成。

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Stage III: Sperm Transfer

As in other scorpions, sperm transfer in Tityus (Atreus) magnimanus is accomplished via a stationary spermatophore that is flagelliform in shape and structure (7). Males of this species are capable of producing their first spermatophore within seven days of maturing molts and can regenerate spermatophores in a very short period, 24 hours (Ross, personal observation).

與其他蝎子一樣，碩螯戾蝎的精子轉(zhuǎn)移是通過一個固定的鞭狀精莢完成的。雄性碩螯戾蝎能夠在蛻皮成熟后的7天內(nèi)產(chǎn)生第一個精莢，并能在很短的時間內(nèi)再生精莢，24小時(來自Rose的個人觀察)。

Once a suitable surface is selected by a male, he pulls the female close and with his chelicerae grasps either the anterior edge of her carapace (n = 10; 83.3%) or chelicerae (n = 2; 16.7%). The courting pair remains still during the release of the spermatophore. The male lowers his venter against the selected surface and the pedicel is attached to the surface. Whereas the spermatophore is slowly extruded, the male gradually elevates his body while simultaneously moving backwards until the trunk and flagellum of the spermatophore are exposed.

一旦雄性選擇了一個合適的表面，它就會把雌性拉近，用它的鉗子抓住它的甲殼的前緣（n = 10;83.3%）或螯肢（n = 2;16.7%）（注：很納悶是怎么抓住的，雌蝎把牙伸出來讓雄蝎抓嗎？……）。在精子被釋放的過程中，這對求偶的蝎子保持靜止不動。雄蝎的腹部向選定的表面降低讓精莢附著在表面上。當(dāng)精莢被緩慢擠出時，雄性逐漸抬起身體，同時向后移動，直到精莢的莖部和鞭毛露出來。

Spermatophore expulsion is marked by the male elevating his entire metasoma vertically and slowly waving it from side-to-side (15 to 39 seconds). Once the spermatophore is deposited, male metasomal movements cease (n = 12; 100%) and he pulls the female toward him and directs her movements until her genital aperture is positioned directly above the erect spermatophore. Once the female genital orifice is positioned above the capsule, the male will rock the female back and forth (6 to 10 seconds) so that the spermatophore is engaged by her genital operculum. During the back-and-forth rocking movements, the female lowers her body upon the trunk of the spermatophore that houses the sperm vesicle and sperm duct, so that the sperm mass is released into her genital aperture. In this observation, as soon as the sperm mass was transferred, females (n = 12; 100%) immediately separated and retreated 4 to 10 cm away from males.

精莢排出的標(biāo)志是雄性將整個后體垂直抬起，慢慢地從一邊到另一邊擺動（15~39秒）。一旦精莢被放置，雄性后體運動停止（n =12;100%），雄性將雌性拉向自己并引導(dǎo)她的動作，直到她的生殖器孔位于豎直的精莢的正上方。一旦雌蟲的生殖孔位于莢膜上方，雄蟲就會前后搖動雌蟲（6~10秒），這樣精莢就會被雌蝎的生殖蓋包裹住。在前后搖擺的過程中，雌性把身體放在裝有精囊和精管的精莢莖部上，這樣精子團就被釋放到她的生殖孔中。在這項觀察中，精子一經(jīng)轉(zhuǎn)移，雌性（n =12;100%）立即分開，并退到離雄性4~10厘米遠(yuǎn)的地方。

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Stage IV: Separation

In all sequences in the present study, mating terminated when the female pulled away and moved a short distance from the male immediately following sperm transfer. As soon as a female started to struggle and pull away, a male simply released his grasp upon female. Once a female moved away from a male, she either rubbed her venter upon the substratum (n = 7; 58.3%) or used one or both pairs of anterior walking legs to vigorously rub her genital operculum (n = 5; 41.7%).

第四階段:分離

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在本研究的所有序列中，當(dāng)雌性在精子轉(zhuǎn)移后立即抽離并離開雄性一小段距離時，交配就終止了。一旦雌性開始掙扎并掙脫，雄性就會松開對雌性的控制。一旦雌性離開雄性，她要么在基底上摩擦她的生殖器（n=7；58.3%）或用一只或一對前腿用力摩擦生殖蓋（n = 5；41.7%）。

The male remained with a spent spermatophore and rapidly moved the tarsi of the first pair of walking legs over the base and trunk of the spermatophore (n = 9: 75%) (4 to 8 seconds) or remained stationary without exhibiting any activity (n = 3; 25%) (30 to 120 seconds).

雄蝎留下一個用過的精莢，在精莢的根部和軀干上快速移動第一對行走腿的跗足（n=9.75%）（4 ~ 8秒），或者靜止不動（n = 3;25%）（30 ~ 120秒）。

After the recuperative stage, males (n = 12; 100%) moved slightly backwards and used their chelicerae to sever the flagellum near its base on the spermatophore. After severing the flagellum, males rested for a brief period (2 to 6 minutes) before becoming vagile and moving away from spermatophores. Consumption of the spermatophore by male or female was not observed in any mating sequence. Post- mating males and females do not exhibit agonistic or aggressive behaviors toward each other and will aggregate in large, mixed gender groups in laboratory enclosures.

休息期過后，雄性（n=12;100%）稍微向后移動，用它們的螯肢切斷位于精莢基部附近的鞭毛。在剪去鞭毛后，雄性會休息一段時間（2至6分鐘），然后開始漫游并離開精莢。在任何交配順序中都沒有觀察到雄性或雌性對精莢的消耗。（注：一直以為是吃精莢，見發(fā)布的視頻）交配后的雄性和雌性不會對彼此表現(xiàn)出敵對或攻擊行為，并且會聚集在實驗容器中的大型混合性別群體中。

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DISCUSSION

The courtship and mating behaviors of Tityus (Atreus) magnimanus are similar to behaviors described in other scorpions including several species within the genus Tityus. However, in order to explain and more thoroughly understand patterns of phylogeny, ecology, biology, reproductive evolution and diversification among scorpions, a precise knowledge of the reproductive biology and associated behaviors in scorpions is required (12).

討論

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碩螯戾蝎的求偶和交配行為與其他蝎子（包括戾蝎屬的幾個物種）的行為相似。然而，為了解釋和更徹底理解蝎子的系統(tǒng)發(fā)育、生態(tài)學(xué)、生物學(xué)、生殖進(jìn)化和多樣化模式，需要對蝎子的生殖生物學(xué)和相關(guān)行為有精確的了解。

The lengthy and time consuming process of the promenade á deux has been traditionally interpreted as the time necessary for a courting male to locate and select a suitable surface upon which he will deposit his spermatophore. However, when a courting male engaged in the promenade á deux is presented with two different surface structures (a stick and a flat rock) during successive mating trials, the male will select the flat rock instead of the stick for spermatophore deposition during one trial and when introduced into another mating arena with a new female, chooses the stick instead of the rock.

這一漫長而耗時的過程傳統(tǒng)上被解釋為求偶的雄性尋找并選擇一個合適的表面來放置他的精莢所需要的時間。然而，在連續(xù)的交配試驗中，對在雙人漫步中的求偶雄性，在連續(xù)的交配試驗中呈現(xiàn)兩種不同的表面結(jié)構(gòu)（枯枝和平坦的巖石）時，它們會在一次試驗中選擇平坦的巖石而不是枯枝來放置精莢，當(dāng)被引入另一個與新雌性交配的場所時，選擇枯枝而不是巖石。

Additionally, a male may direct the female over a surface dozens of times before finally selecting the surface as the site of spermatophore deposition. Francke (7) reported that males of Centruroides sculpturatus Ewing, 1928 (Buthidae) when maintained overnight in plastic bags with pieces of crumpled newsprint for cover would deposit spermatophores upon the newsprint.

此外，雄性在最終選擇一個表面作為精莢放置的地點之前，可能會引導(dǎo)雌性在一個表面上經(jīng)過幾十次。Francke曾報道，雄性雕紋似刺尾蝎放在塑料袋中過夜時，里面放一塊皺報紙蓋住，它會將精莢放置在報紙上。

Overall, a courting male does not appear to be overly selective in his choice of site for spermatophore deposition. Personal observations of additional mating sequences in Tityus (Atreus) magnimanus and Tityus (Atreus) trinitatis Pocock, 1897 have revealed that a courting male will select any solid surface encountered for spermatophore deposition including compacted substrate, fragments of clay plant pots, pieces of peat seedling pots, rocks, sticks, various tree barks, dried leafs, sections of palm root, quarter dollars, plastic bottle caps, cardboard and construction paper.

總的來說，求愛的雄性在選擇精莢放置的地點時似乎沒有過度的選擇性。通過個人對碩螯戾蝎和特立尼達(dá)戾蝎的額外交配序列觀察發(fā)現(xiàn)，求偶的雄性會選擇任何遇到的固體表面來放置精莢，包括壓實的基質(zhì)、粘土花盆的碎片、泥炭苗盆的碎片、巖石、樹枝、各種樹皮、干葉子、棕櫚根的部分、25美分硬幣、塑料瓶蓋、紙板和建筑紙。

Male juddering has been reported to occur in 18 species, from eight families, and has been reported to occur during all stages of courtship (3, 4). Juddering behavior during the promenade á deux was observed in ten pairings of Tityus (Atreus) magnimanus and was utilized by a male in response to a female that became unresponsive to male-directed movements or attempted to move in an opposite direction of male during courtship. In two pairings, females did not resist male- directed movements or attempt to pull away and move in an opposite direction. In both pairings, juddering was not displayed by either male during any stage of courtship. These observations suggest that juddering by male Tityus (Atreus) magnimanus may be used in direct response to resistance behaviors exhibited by females during the promenade and may act to stimulate mating.

據(jù)報道，雄性抖動發(fā)生在8個科的18個物種中，并且發(fā)生在求偶的所有階段。在10對碩螯戾蝎中觀察到在雙人漫步期間的抖動行為，這是雄性對雌性在求偶期間對雄性引導(dǎo)的動作沒有反應(yīng)或試圖向相反方向移動的反應(yīng)。在兩對交配蝎子中，雌性沒有抵抗雄性的動作，也沒有試圖抽離并朝相反的方向移動。這兩對中的雄性在求偶的任何階段都沒有表現(xiàn)出抖動。這些觀察結(jié)果表明，雄性碩螯戾蝎的抖動可能是對雌性在散步時表現(xiàn)出的抵抗行為的直接反應(yīng)，也可能是刺激交配的行為。

Juddering by the male during the initial stage did not occur in any of the 12 pairings observed during this study. Males introduced into mating arenas containing females either sensed their movements and moved directly to them or encountered immobile females during random movements. Males did not exhibit pre-contact mate recognition behaviors and readily approached moving and immobile females. Once contact was made with a female, the male would immediately grasp a single pedipalp of the female and make repeated grasps upon the various segments. In two pairings, the male made repeated grasps upon metasomal segments III to V of the female.

在這項研究中觀察到的12對配對中，雄性在初始階段沒有出現(xiàn)抖動。被引入有雌性的交配場所的雄性要么感覺到它們的動作并直接向它們移動，要么在隨機移動中遇到不動的雌性。雄性沒有表現(xiàn)出接觸前的配偶識別行為，并且很容易接近活動和不活動的雌性。一旦與雌性接觸，雄性會立即抓住雌性的一條須肢，并反復(fù)抓住不同的部分。在兩對配對中，雄性反復(fù)抓住雌性的第III到V節(jié)后體。

Alexander (1) noted that there is a marked sexual dimorphism of the metasomal segments in males and females of Tityus (Atreus) trinitatis and that courting males would approach females and repeatedly grasp various metasomal segments. Furthermore, the author suggested that males utilize grasping behaviors to identify the sexually dimorphic characters of the metasomal segments in females. The exhibition of grasping behaviors in courting males of Tityus (Atreus) magnimanus and Tityus (Atreus) trinitatis propose that differences in the morphology of various structures (chelae, pedipalps, metasomal segments) are important for mate recognition by males of both species during courtship.

Alexander注意到特立尼達(dá)戾蝎的雄性和雌性之間存在著明顯的兩性二態(tài)性，求偶的雄性會接近雌性并反復(fù)抓住后體的不同節(jié)。此外，他表示，雄性利用抓握行為來識別雌性后體關(guān)節(jié)的兩性二態(tài)特征。碩螯戾蝎和特立尼達(dá)戾蝎在求偶過程中表現(xiàn)出的抓握行為表明，兩種雄性在求偶過程中各種結(jié)構(gòu)（螯、須肢、后體關(guān)節(jié)）的形態(tài)差異對它們的擇偶識別很重要。