Hoarding of acorns by granivorous mice and its role in the population processes of Pasania edulis (Makino) Makino

In order to evaluate the role of granivorous mice, Apodemus speciosus Temminck and Apodemus argenteus Temminck, in the regeneration of Pasania edulis (Makino) Makino, an evergreen tree, we conducted field studies examining acorn crops, the population dynamics and hoarding behavior of mice and pre- and postdispersal acorn predation in an evergreen broad-leaved forest dominated by P.edulis in Kagoshima, southern Japan. The study was conducted from 1994 to 2000. Apodemus mice selected sound acorns and hoarded them in the soil after transporting them for distances ranging from 5.4 to 19.9m. This behavior appeared to be beneficial to the establishment of seedlings of P.edulis. However, the positive effects were negated by a high rate of recovery of the acorns hoarded by the mice. A life-table analysis of five cohorts of P.edulis indicated that postdispersal predation of acorns by Apodemus mice was a major contributor to annual variation in the number of new seedlings. A considerable number of acorns germinated only in years with a good acorn crop and a low level of resident mice. These results suggest that Apodemus mice are serious acorn predators rather than important dispersers. After germination, acorns were attacked by the acorn borer at a high rate. From these results, we can conclude that acorn production probably makes a smaller contribution to the regeneration of P.edulis than coppice shoot production at the study site.     

昆虫寄生对辽东栎种子命运的影响

报道了昆虫寄生对辽东栎 Quercus liaotungensis 种子命运的影响。结果表明：（1）被昆虫寄生的种子发芽率明显低于正常成熟的种子,被昆虫寄生的种子或者不能发芽,或者能发芽但随后死亡;（2）从虫卵到老龄幼虫的发育在橡实内完成,当橡实落地后幼虫破壳而出,整个种子库的幼虫破壳时间延续2～50天,并非高度同步化;种子库的虫寄生率高达45.4%,以柞栎象Curculio dentipes为主,还有少量其它种类,如另一种象虫Curculio sp.、栗白小卷蛾 Cydia kurokoi 和螟蛾科（Pyralidae）的一未定名种;（3）昆虫寄生影响橡实成熟,测量种子大小分布表明,被昆虫寄生的种子的大小分布介于未成熟和成熟种子之间;（4）象虫寄生取食引起16.05% 的种子能量损失,显著低于对照组。结果证明昆虫寄生对辽东栎种子的命运有重要影响,是影响辽东栎种群更新的关键因素之一。     

都江堰地区灌丛内袍栎橡子的捕食和搬运

从2000年9月至2002年12月,以四川都江堰常绿阔叶林灌丛内的优势种--枹栎(Quercus serrata)为研究对象,研究了昆虫、鼠类和真菌对枹栎橡子的捕食和搬运.结果表明(1)取食橡子的鼠类的群落结构和数量在年间差异较小,这可能与三年间袍栎橡子的产量相对稳定有关;(2)袍栎橡子的虫蛀率在三年间存在显著性差异,范围在14.17%-32.49%之间;(3)鼠类的就地取食和搬运是影响种子库内橡子命运的主要因素;(4)橡子虫蛀率的变化影响鼠类的觅食活动.鼠类对饱满橡子和虫蛀橡子的选择性取食和搬运可能影响许多坚果植物(栎树)的自然更新过程[动物学报50(4)535-540,2004].     

都江堰地区灌丛内迺栎橡子的捕食和搬运(英文)

从 2 0 0 0年 9月至 2 0 0 2年 1 2月 ,以四川都江堰常绿阔叶林灌丛内的优势种———栎 (Quercusserrata)为研究对象 ,研究了昆虫、鼠类和真菌对栎橡子的捕食和搬运。结果表明 :(1 )取食橡子的鼠类的群落结构和数量在年间差异较小 ,这可能与三年间栎橡子的产量相对稳定有关 ;(2 )栎橡子的虫蛀率在三年间存在显著性差异 ,范围在 1 4 1 7% - 32 4 9%之间 ;(3)鼠类的就地取食和搬运是影响种子库内橡子命运的主要因素 ;(4 )橡子虫蛀率的变化影响鼠类的觅食活动。鼠类对饱满橡子和虫蛀橡子的选择性取食和搬运可能影响许多坚果植物 (栎树 )的自然更新过程     

啮齿动物鉴别虫蛀种子的能力及其对坚果植物更新的潜在影响

作为种子捕食者和种子扩散者, 啮齿动物对产坚果植物的自然更新有很大作用。然而, 对啮齿动物鉴别虫蛀种子的能力颇有争议。2001 年秋季在中国四川都江堰市实验林场, 以3 种比例(I₁∶S = 1∶1 , I₁∶S = 4∶1 和I₁∶I₂∶S = 1∶1∶1)供给啮齿动物4 种壳斗科种子: 栓皮栎( Quercus variabilis) 、枹树( Q. serrata) 、青冈( Cyclobalanopsis glauca) 和栲树( Castanopsis fargesii) 的3 种坚果型(即饱满种子(S) , 不含象虫的虫蛀种子(I₁) 和含象虫的虫蛀种子( I₂ ) 验证了啮齿动物能够准确地鉴别虫蛀种子这一假说。结果表明, 在3 种比例下4 种坚果的虫蛀种子的消失速率均慢于饱满种子。即使虫蛀种子的比例增加, 啮齿动物显著地搬走了更多的饱满种子(67%～92%) 。当虫蛀种子的比例增加时, 虫蛀种子就地消耗和拒绝的比例降低, 搬走比例增加。啮齿动物并不拒绝虫蛀种子, 这可能与它们的可利用性(如象虫可以作为蛋白质的补充) 和数量以及它们的觅食行为有关。结果证实啮齿动物能准确地鉴别虫蛀种子, 从而有区别地搬走并贮藏更多的饱满种子, 消耗一部分可以利用的虫蛀种子(包括其内的象虫) 。这样, 啮齿动物通过对饱满种子和虫蛀种子的鉴别和选择会影响不同种子的命运, 从而对这些坚果植物的自然更新产生影响。     

Large acorns benefit seedling recruitment by satiating weevil larvae in <Emphasis Type="Italic">Quercus aliena</Emphasis>

Plants can enhance their seed viability and survival through reducing the fitness costs of herbivory by satiating seed predators at either plant or seed level, or both. The former satiates with production of large crops while the latter with large-sized seeds, enhancing the probability of seed survival. We analyzed both types of satiation in the interaction between the Quercus aliena and the weevil Curculio spp in 2 years with different crop size and acorn size. Larger crop size seemed to satiate less effectively weevils due to higher proportions of the seeds were attacked in oaks producing more crops of smaller acorns. However, prematurely abscission of infested acorns in large crop year might compensate this ineffectiveness. Larger acorns in lean year well satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival and seedling establishment. Although effective satiation by larger acorns would be negated by smaller seed crops, the proportion of attacked acorns that survived increased with large acorn size in lean year. These results highlighted the importance of annual variation in acorn size of Quercus aliena in satiating and defending weevil larval predations, consequently in successful seedling recruitment and establishment.     

Sucrose in storage media and cultivar affects post-storage regrowth of <Emphasis Type="Italic">in vitro</Emphasis> Hosta propagules

The goal of this research was to investigate if culturing in high sucrose (5%) liquid media during multiplication phase (stage II) would enhance endogenous sugar levels and dry matter sufficiently to allow storage of in vitro plants in sugar free media without adversely affecting post-storage recovery. Hosta tokudama Newberry Gold (NBG) and Hosta Striptease were cultured in Murashige and Skoog (MS) media containing 5% sucrose during stage II and transferred to rooting phase (stage III) in MS medium without (0%) sucrose or with 3% sucrose for 4weeks. At the end of stage III, cultures were stored, with the remaining media, at 10°C with 5molm^–2s^–1 photosynthetic photon flux (PPF) from cool white fluorescent lamps for 7 or 14weeks with or without a 2-week dark period prior to removal from storage. In both cultivars, stage III plants cultured in 3% sucrose media had higher soluble sugar levels and greater shoot and root biomass than those cultured in 0% sucrose media. Shoot and root soluble sugars decreased during storage. Shoot growth ceased during storage in both media. Root dry matter continued to increase in plants stored in 3% sucrose media but did not change in 0% sucrose media. Plants cultured in 3% sucrose media had less leaf chlorosis and less mortality after 7 or 14weeks of low temperature storage than the plantlets from sugar free media. Extending the storage period from 7 to 14weeks or introduction of 2-week dark period at the end of storage did not affect leaf chlorosis or plant mortality during acclimatization. Post-storage growth varied with the cultivar. Benefit of having sucrose in storage media was to develop a strong root system that aided the acclimatization and post-storage growth following 7 or 14week storage. Sucrose loading by culturing plants in liquid media containing 5% sucrose did not allow storage in sugar free media without adversely affecting post-storage growth in both cultivars.     

Five-year study on the autumn food habits of the Asiatic black bear in relation to nut production

We studied the autumn food habits of the Asiatic black bear (Ursus thibetanus) from 1993 to 1997, based on 202 fecal samples in the Chichibu Mountains, central Japan. Nuts occupied the highest proportions of autumn foods (59.9–85.8% important values). Although the proportion of nuts of Quercus crispula, Fagus crenata and Fagus japonica varied greatly between the years, acorns of Q.crispula were most prevalent in four of the five years. We also determined the relative nut production of these three species by counting the number of nuts or cupules on the ground. Black bears consumed the nuts according to their relative availabilities. Nuts of Q.crispula appeared to be the most important food because: (i) these nuts were eaten in the highest proportion in four of the five study years; and (ii) even in poor years the bears consumed acorns of Q.crispula, whereas nuts of Fagus spp. were not consumed. We discuss the significance of alternative foods for black bears in relation to food tree diversity in the forest and the necessity for long-term studies examining the food habits of Asiatic black bears.     

Prolonged diapause and seed predation by the acorn weevil,Curculio robustus,in relation to masting of the deciduous oak Quercus acutissima

Many researchers have studied the relationship between masting by trees and seed predation by insects. Most of these studies have been plant centered, with little focus on the insect perspective. To estimate the effect of mast seeding on insect seed predators, the life‐history traits of these insects must also be considered because some seed insects can survive lean years by prolonged diapause. In this study, I examined larval infestation of acorns and life‐history traits of the acorn weevil, Curculio robustus (Roelofs) (Coleoptera: Curculionidae), in relation to acorn production of the deciduous oak Quercus acutissima Carruthers (Fagaceae) in a coppice stand in central Japan in 2004–2009. Curculio robustus females oviposit into Q. acutissima acorns, inside which the larvae develop. Mature larvae leave acorns and burrow into the soil, where they overwinter. Although germination did occur in acorns infested by weevil larvae, the percentage of germination was lower in acorns damaged by many larvae. Acorn production in Q. acutissima varied considerably among years. Both the number of C. robustus larvae infesting acorns and the percentage of acorns infested were affected by the amount of acorns produced by Q. acutissima, and two successive lean years appeared to have a considerable impact on C. robustus population size. Consequently, only a small fraction of the acorns produced were lost to predation in a mast year after two successive lean years. However, C. robustus could survive the two successive lean years because of prolonged larval diapause, probably leading to a marked decrease in population size. These findings suggest that masting in Q. acutissima succeeds as a predator satiation strategy in response to acorn damage by C. robustus, and that C. robustus has developed prolonged diapause as a counter‐adaptation.     

Early development of the self-fertilizing mangrove killifish <Emphasis Type="Italic">Rivulus marmoratus</Emphasis> reared in the laboratory

The mangrove killifish Rivulus marmoratus was reared at 25°±1°C and 17ppt salinity from 0 to 100 days after hatching (DAH), and its early development was described by examining growth and morphometric parameters, meristic characters (vertebral and fin-ray counts), bone-cartilage development, and pigmentation. Growth was isometric for preanal length, head length, snout length, body depth, pectoral-fin length, dorsal-fin length, anal-fin length, and caudal-peduncle depth. Negative allometric growth was observed in eye diameter and gape size. Meristic counts (mean±SD) for vertebrae (34.2±0.4) and dorsal- (8.6±0.5), anal- (11.4±0.5), and caudal-fin rays (30.2±0.8) were complete at 0 DAH (n=5), whereas pectoral-fin rays and pelvic-fin rays were complete by 30 DAH (14.5±0.4, n=5) and 60 DAH (4.2±0.8, n=5). Full ossification of meristic elements proceeded in the following sequence: vertebrae (by 30 DAH), caudal-, dorsal-, and anal-fin rays (by 60 DAH), pectoral-fin rays (between 60 DAH and 100 DAH), and pelvic-fin rays (by 100 DAH). Both morphological characters and meristic counts indicate that this species can be considered to be a juvenile after 9.8mm in standard length (20 DAH).     

Fine root biomass,production and turnover in a fertilized <Emphasis Type="Italic">Larix leptolepis</Emphasis> plantation in central Korea

The effects of fertilization [control (C), 200kgNha^–1+25kgP ha^–1 (LNP) and 400kgNha^–1+ 50kgP ha^–1 (HNP)] on fine root dynamics were examined in a 40-year-old Larix leptolepis plantation in central Korea. The average fine root biomass during the growing season for C, LNP and HNP was 957, 934 and 814kgha^–1, respectively, whereas the fine root production for C, LNP and HNP was 2103, 2131 and 2066kgha^–1, respectively. Nitrogen and P inputs into the soil via fine root turnover for C, LNP and HNP were 23.0 and 1.2, 23.3 and 1.2 and 22.6 and 1.2kgha^–1, respectively. There were no significant differences in fine root biomass, production and N and P inputs through fine root turnover between the fertilization treatments during the first growing season after fertilization.     

Consequences of harvesting for genetic diversity in American ginseng (<Emphasis Type="Italic">Panax quinquefolius</Emphasis> L.): a simulation study

American ginseng, Panax quinquefolius L., is one of the most heavily traded medicinal plants in North America. The effect of harvest on genetic diversity in ginseng was measured with a single generation culling simulation program. Culling scenarios included random harvest at varying levels, legal limit random harvest and legal limit mature plant harvest. The legal limit was determined by the proportion of legally harvestable plants per population (% mature plants per population). Random harvest at varying levels resulted in significant loss of genetic diversity, especially allelic richness. Relative to initial levels, average within-population genetic diversity (H _e) was significantly lower when plants were culled randomly at the legal limit (Mann–Whitney U=430, p<0.001) or when only mature plants were culled (Mann–Whitney U=394, p<0.01). Within-population genetic diversity was significantly higher with legal limit mature plant harvest (H _e=0.068) than when plants were culled randomly at the legal limit (H _e=0.064; U=202, p<0.01). Based on these simulations of harvest over one generation, we recommend that harvesting fewer than the proportion of mature plants could reduce the negative genetic effects of harvest on ginseng populations.     

Population density and group composition of Japanese sika deer (<Emphasis Type="Italic">Cervus nippon yakushimae</Emphasis>) in an evergreen broad-leaved forest in Yakushima,southern Japan

The population density of Japanese sika deer (Cervus nippon yakushimae Kuroda and Okada) in an evergreen broad-leaved forest in Yakushima, southern Japan, was surveyed over 4years from 1998 to 2001. Two approximately 50ha study sites, Hanyama and Kawahara, were established with a total of 4km of census trails at each site. The estimated densities of sika deer at the two sites were 43–70 deerkm^–2 at Hanyama and 63–78 deerkm^–2 at Kawahara, although these values might be underestimates. The adult sex ratio (number of adult males:number of adult females) ranged from 0.6 to 1.0 at Hanyama, and from 0.4 to 0.9 at Kawahara. Mean group size was 1.9 deer (male group, 1.5 deer; female group, 1.6 deer; mixed group, 3.6 deer). The population density of sika deer was relatively high compared to other sites in Japan, with the exception of very small (<10km²) islands. Possible explanations for this naturally high density of sika deer in an evergreen broad-leaved forest in Yakushima are discussed.     

Alarm response of hatchlings of the apple snail, <Emphasis Type="Italic">Pomacea canaliculata</Emphasis> (Gastropoda: Ampullariidae), to aqueous extracts of other individuals

We examined how hatchlings of the freshwater snail, Pomacea canaliculata, responded to aqueous extracts of conspecific hatchlings. Three, 3-day-old hatchlings were macerated in deionized water (1mg hatchling per 1ml water). When 0.5ml of the aqueous extract was added to a test tube containing 10 hatchlings of the same age and 50ml of water, the hatchlings in the water began to crawl out of the water within 5min. The proportion of hatchlings that crawled out of the water approached 0.6–0.9 after 1h, but gradually decreased to 0.4 after 24h. The relatedness between the live and the macerated hatchlings had no significant influence on the response. Hatchlings of egg masses obtained either in the laboratory or in the wild responded similarly to aqueous extracts of hatchlings from either egg mass. This suggests that the conditions under which the egg masses were incubated or the conditions that their parents had experienced had no effect on the hatchlings response. When compared with experiments reported on other aquatic animals, we consider the behavior of the hatchlings to be an alarm response of escaping from predators.     

Larvae of <Emphasis Type="Italic">Arnoglossus elongatus</Emphasis> (Pleuronectiformes: Bothidae) from northwestern Australia

Eleven postflexion larvae (9.1–23.9mm standard length: SL) of a bothid, Arnoglossus elongatus, from northwestern Australian waters were described. These specimens were characterized by vertebral numbers of 11–12+32–34=43–45, slender body, remarkably elongated second dorsal fin ray, and presence of melanophores on and slightly above midlateral line of body, plus on dorsal wall of abdominal cavity and air bladder (specimens >12.6mm SL), proximally on caudal fin rays (>16.5mm SL), and on pterygiophore zones of dorsal fin and anal fin (>23.8mm SL).     

<Emphasis Type="Italic">In vitro</Emphasis> screening of sugarcane to evaluate smut susceptibility

Tissue-cultured plantlets of three sugarcane (Saccharum spp.) cultivars having a known field smut reaction were screened for susceptibility to Ustilago scitaminea H&P Sydow. Plantlets were inoculated with 0.5 l of a suspension of equally mixed quantities of plus and minus mating type sporidia of U. scitaminea at concentrations ranging from 1×10¹ to 1×10⁶ cells. Fungal sori (whips) were produced in cultivar N12 (intermediate) 6weeks following inoculation with 1×10⁵ mixed sporidia and thereafter in cultivar NCo310 (susceptible) but not in cultivar N19 (resistant). Sori bearing teliospores were produced up to 3months following inoculation and incubation at 26°C. No sori were produced at mixed sporidial concentrations lower than 1×10⁵cells. The in vitro soral production in cultivars N19, N12 and NCo310 was 0, 27.5 and 47.5% respectively. Plantlets inoculated with 1×105sporidia of only one mating-type did not produce sori in any of the three cultivars tested. Blind scoring of an unknown sugarcane cultivar by this method corresponded exactly with its field smut rating.     

Carbon dioxide and water vapor exchange over a <Emphasis Type="Italic">Miscanthus</Emphasis>-type grassland: Effects of development of the canopy

The eddy correlation technique was employed to measure net ecosystem carbon dioxide (CO₂) (NEE) and water vapor exchange (LE) over a C3/C4 co-occurring wet temperate Miscanthus-type grassland in the Kanto plain of Japan in the 1999 growing season. The maximal mean canopy height and maximal leaf area index were 1.0m and 5.5, respectively. The daily maximal LE was approximately 540Wm^–2. The maximum value of daily accumulative LE was 16.3MJday^–1. Daily variation of the decoupling factor () suggests that in the morning LE decoupled with the atmosphere, and the available energy was the major driving force for LE, whereas in the afternoon LE coupled strongly with the atmosphere, and the atmospheric evaporative demand played a critical role in LE. The decline in (from 0.8 to 0.5) with the growing season demonstrates that LE decoupled from the atmosphere in the later growth season. The peak NEE value was 57.4µmolCO₂m^–2s^–1 (the positive value signifies the canopy carbon gain was from the air). The maximal daily integrated NEE was 1.06molCO₂m^–2day^–1 observed during the peak growth stage. A rectangular hyperbolic model was used to describe the relation between daytime NEE and incident photosynthetic photon flux density (PPFD). The net ecosystem CO₂ was not light-saturated up to a PPFD level of 2000µmol m^–2s^–1. The initial slope estimated with the NEE–PPFD response model was approximately 0.042molCO₂mol^–1photon on average. The canopy light compensation point ranged from 210 to 430µmolm^–2s^–1 with an average of approximately 310µmolm^–2s^–1. Both the initial slope and the canopy light compensation point decreased as the canopy senesced. The switch in dominance from C3 to C4 plants played an important role in the canopy fluxes.     

Multi-trophic effects of ungulate intraguild predation on acorn weevils

Predators and parasitoids may contribute to controlling the population sizes of phytophagous insects, and this has been shown to benefit plants. Phytophagous insects may also be killed by other herbivores (intraguild predation), usually larger-sized vertebrates that ingest insects accidentally while feeding on common food sources. We studied the intraguild predation on acorn weevils by ungulates and assessed the consequences for weevil populations. Infested acorns are prematurely abscised and the weevil larvae finish their development inside the acorns after being dropped. Our results show that weevil larvae were killed by ungulates eating the infested acorns on the ground. Ungulates did not discriminate between infested and sound acorns, and the probability of a larva being incidentally eaten was inversely related to acorn availability. Thus, predation risk was enhanced by the premature drop of infested acorns when acorn availability on the ground was low. Predation rates on infested acorns were much higher where ungulates were present, and acorn infestation rates were significantly lower. However, ungulates did not provide the oaks any net benefit, since the reduction of infestation rates was not enough to compensate for the large amounts of sound acorns eaten by ungulates. Seed predation is usually studied as a progressive loss of seeds by pre- and post-dispersal predators, but the interactions between them are usually not considered. We show that intraguild predation on insects by large ungulates had an effect on the structure of the foraging guild, as the proportion of acorns predated by insects decreased; however, replicating the same experimental design in different ecological scenarios would increase the strength of these results. In conclusion, the present study shows the importance of considering the multi-trophic interactions between seed predators in order to have a complete picture of granivory.     

Ontogeny of the mandarinfish <Emphasis Type="Italic">Siniperca chuatsi</Emphasis> (Perciformes: Sinipercidae) reared in aquarium

Early-stage morphologies of the mandarinfish Siniperca chuatsi are described on the basis of an ontogenetic series of reared specimens in an aquarium. Spherical eggs (diameter 1.70–1.82mm) with a single oil globule (0.40–0.48mm) were free-demersal and easily floated when agitated. Hatching occurred about 3 days after spawning at about 24°C. Newly hatched larvae (3.8–4.2mm in notochord length: NL) had many melanophores on the yolk sac. After reaching ca. 5.5mm NL (8–9+19–20=27–28 myomeres), larvae had almost completely absorbed the yolk, possessed a large mouth and sharp teeth, and were starting to prey on other fish larvae. Three large preopercle spines appeared at ca. 5.5mm NL, five spines by ca. 8.5mm NL, and eight by ca. 21mm in standard length (SL). The interopercle bore a single spine at ca. 8.5mm NL and two spines at ca. 13.5mm SL. A single spine appeared at the supracleithrum and another at the opercle at ca. 10mm SL. Dorsal fin spines and pelvic, anal, and caudal fins were completed at ca. 10mm SL. Dorsal fin rays and pectoral fins were completed at ca. 13.5mm SL. Four ontogenetic characters (free-demersal eggs, large jaws with large teeth, conspicuous head spination, and precocious completion of dorsal fin spines) are rare among freshwater percoids.     

Competition between <Emphasis Type="Italic">Eucalyptus victrix</Emphasis> seedlings and grass species

Competition in a natural system may be interspecific or intraspecific. In semiarid ecosystems, competition for resources between established neighboring grass species and newly recruited seedlings is very high. To examine the effects of grass species density, growing space and time of establishment on Eucalyptus victrix seedlings (interspecific competition), and the effect of density and growing space within E.victrix (intraspecific competition) we conducted an experiment under controlled conditions. We tested four hypotheses (i) E.victrix seedling growth is not affected by grass density; (ii) there is no difference in E.victrix survival and growth between early and later grass establishment; (iii) interspecific competition is not more intense than intraspecific competition in E.victrix; and (iv) growth of E.victrix seedlings is not dependent on available growing space. In a monoculture of E.victrix, seedling mortality was higher (10%) in large pots. In mixed culture pots, where E.victrix seedlings and grass seedlings were planted on the same day, E.victrix seedlings survived for up to 4weeks, but started to die after week five in the smallest pots. However, mortalities occurred in pots of all sizes when grass was established before E.victrix seedlings. Results also indicated that the resources necessary for the growth of individual E.victrix seedlings were more limiting under conditions of increased density of neighboring grass species rather than intraspecific competition. In particular, photosynthetic area of E.victrix seedlings was drastically reduced in mixed cultures. Although density, pot size and time of planting had impacts on E.victrix seedlings, the patterns of these impacts were variable.