跟著城市嚮導「老臺北胃」，用味道認識臺北

很多朋友來臺北，
都會問我同一個問題：
「臺北小吃那麼多，到底該從哪裡開始吃？」
夜市裡攤位一字排開、老店藏在巷弄轉角，
看起來都很有名，卻又怕吃錯、踩雷，
結果行程走完，反而沒真正記住臺北的味道。
我常被朋友笑說是「老臺北胃」。
不是因為特別會吃，而是因為在這座城市待久了，
知道哪些味道是陪著臺北人成長的日常。
這篇文章，就是我整理的一份清單。
如果你第一次來臺北，
我會帶你從這 10 樣最具代表性的臺北小吃開始，
不追一時爆紅、不走浮誇路線，
而是讓你吃完後能真正理解
原來，這就是臺灣的小吃文化。
跟著老臺北胃走，
用最簡單的方式，
把臺北的味道，一樣一樣記在心裡。

我怎麼選出這 10 大臺北小吃？

在臺北，
你隨便走進一條夜市或老街，
都可以輕易列出 30 種以上的小吃。
所以這份清單，
不是「臺北最好吃」的排名，
 而是我站在「第一次來臺北的旅客」角度，
做的推薦。
身為一個被朋友稱作「老臺北胃」的人，
我選這 10 樣小吃時，心裡一直放著幾個原則。

一吃就知道：這就是臺灣味

燒烤、火鍋很好吃，
但換個城市、換個國家，也吃得到。
我挑的，是那種
只要一入口，就會讓人聯想到的臺灣味。
 不需要解釋太多，舌頭就能懂。

不只是好吃，而是有「臺北日常感」

臺北的小吃迷人，
不只在味道，
而在它融入生活的方式。
我在意的是：

1. 會不會出現在早餐、宵夜、下班後
2. 有沒有陪伴這座城市很久的記憶

吃完之後，你會記得臺北

最後一個標準很簡單。
如果你回到家，
還會突然想起某個味道、某碗熱湯、某個攤位的香氣
那它就值得被放進這份清單裡。

接下來的 10 樣臺北小吃，
就是我會親自帶朋友去吃的在地美食。
不趕行程、不拚數量，
而是一口一口，
慢慢認識臺北。

第 1 家：饌堂－黑金滷肉飯（雙連店）｜一碗就懂臺灣人的日常

如果只能用一道料理，
 來解釋臺灣人的日常飲食，
 那我一定會先帶你吃滷肉飯。
在臺北，滷肉飯不是什麼特別的節慶料理，
 而是從早餐、午餐到宵夜，
 默默陪著很多人長大的味道。
而在眾多滷肉飯之中，
饌堂－黑金滷肉飯（雙連店），
 我很常帶第一次來臺北的朋友造訪的一家。

為什麼第一站，我會選饌堂？
饌堂的滷肉飯，走的是**「黑金系」路線**。
滷汁顏色深、香氣厚，
卻不死鹹、不油膩。
滷肉切得細緻，
肥肉入口即化，搭配熱騰騰的白飯，
每一口都是很完整、很臺灣的味道。
對第一次吃滷肉飯的旅客來說，
這種風味夠經典、也夠穩定，
不需要太多心理準備，就能理解為什麼臺灣人這麼愛它。

不只是好吃，而是「現在的臺北感」
饌堂並不是那種躲在深巷裡的老攤，
空間乾淨、節奏俐落，
卻沒有失去滷肉飯該有的靈魂。
這也是我會推薦給旅客的原因之一：
它保留了臺灣小吃的核心味道，
同時也讓第一次來臺北的人，
吃得安心、坐得舒服。

老臺北胃的帶路小提醒
如果是第一次來：

1. 一定要點招牌黑金滷肉飯
   
2. 可以加一顆滷蛋，風味會更完整
3. 搭配簡單的小菜，就很有臺灣家常感

這不是那種吃完會驚呼「哇！」的料理，
而是會讓你在幾口之後，
慢慢理解
原來，臺灣人的日常，就是這樣被一碗飯照顧著。

地址：103臺北市大同區雙連街55號1樓

電話：0225501379

菜單：https://bio.site/ZhuanTang

第 2 家：富宏牛肉麵｜臺北深夜也醒著的一碗熱湯

如果說滷肉飯代表的是臺灣人的日常，
 那牛肉麵，
 就是很多臺北人心中最有份量的一餐。
而在臺北提到牛肉麵，
 富宏牛肉麵，
 幾乎是夜貓族、加班族、外地旅客一定會被帶去的一站。

為什麼老臺北胃會帶你來吃富宏？
富宏最讓人印象深刻的，
不是華麗裝潢，
而是那鍋永遠冒著熱氣的紅燒湯頭。
湯色濃而不混，
帶著牛骨與醬香慢慢熬出的厚度，
喝起來溫潤、不刺激，
卻會在嘴裡留下很深的記憶點。
牛肉給得大方，
燉到軟嫩卻不鬆散，
搭配彈性十足的麵條，
每一口都很直接、很臺北。

不分時間，任何時候都適合的一碗麵
富宏牛肉麵最迷人的地方，
在於它陪伴了無數個臺北的夜晚。
不管是深夜下班、看完演唱會、
或是剛抵達臺北、還沒適應時差，
這裡總有一碗熱湯在等你。
對旅客來說，
這種不用算時間、不用擔心打烊的安心感，
本身就是一種臺北特色。

老臺北胃的帶路小提醒
第一次來富宏，我會這樣點：

1. 紅燒牛肉麵是首選
   
2. 如果想吃得更過癮，可以加點牛筋或牛肚
3. 湯先喝一口原味，再視情況調整辣度

這不是精緻料理，
卻是一碗能在任何時刻撐住你的牛肉麵。
在臺北，
很多夜晚，
就是靠這樣一碗熱湯走過來的。

地址：108臺北市萬華區洛陽街67號

電話：0223713028

菜單：https://www.facebook.com/pages/富宏牛肉麵-原建宏牛肉麵/

第 3 家：士林夜市・吉彖皮蛋涼麵｜臺北夏天最有記憶點的一口清爽

如果你在夏天來到臺北，
 一定會很快發現一件事
 這座城市，真的很熱。
也正因為這樣，
 臺北的小吃世界裡，
 才會出現像「涼麵」這樣的存在。
而在士林夜市，
 吉彖皮蛋涼麵，
 就是我很常帶旅客來吃的一家。

為什麼在夜市，我會帶你吃涼麵？
很多人對夜市的印象，
都是炸物、熱湯、重口味。
但真正的臺北夜市，
其實也很懂得照顧人的胃。
吉彖的涼麵，
冰涼的麵條拌上濃郁芝麻醬，
再加上切得細緻的皮蛋，
入口的第一瞬間，
就是一種「被降溫」的感覺。
那種清爽，
不是沒味道，
而是在濃香與清涼之間取得剛剛好的平衡。

皮蛋，是靈魂，也是臺灣味的關鍵
對很多外國旅客來說，
皮蛋是既好奇、又有點猶豫的存在。
但我常說，
如果要嘗試皮蛋，
涼麵是一個非常溫柔的起點。
芝麻醬的香氣會先接住味蕾，
皮蛋的風味則在後段慢慢出現，
不衝、不嗆，
反而多了一層深度。
很多人吃完後，
都會露出那種「原來是這樣啊」的表情。

老臺北胃的帶路小提醒
第一次點吉彖皮蛋涼麵，我會建議：

1. 一定要選皮蛋款，才吃得到特色
2. 醬料先拌勻，再吃，風味會更完整
3. 如果天氣真的很熱，這一碗會救你一整晚

這不是華麗的小吃，
卻非常臺北。
在悶熱的夜晚，
站在夜市人潮裡，
吃著一碗涼麵，
你會突然明白——

原來臺北的小吃，連氣候都一起考慮進去了。

地址：111臺北市士林區基河路114號

電話：0981014155

菜單：https://www.facebook.com/profile.php?id=100064238763064

第 4 家：胖老闆誠意肉粥｜臺北人深夜最踏實的一碗粥

如果你問我，
 臺北人在深夜、下班後，
 最容易感到被安慰的食物是什麼——
 我會毫不猶豫地說：肉粥。
而提到肉粥，
 胖老闆誠意肉粥，
 就是很多老臺北人口中的那一味。

為什麼這一碗粥，會被叫做「誠意」？
胖老闆的肉粥，看起來很簡單。
白粥、肉燥、配菜，
沒有華麗擺盤，也沒有複雜作法。
但真正坐下來吃，你會發現：
這碗粥，不敷衍任何一個細節。
粥體滑順、不稀薄，
肉燥香而不膩，
搭配各式家常小菜，
一口一口吃下去，
很自然就會放慢速度。
這種味道，
不是要你驚艷，
而是要你安心。

這不是觀光小吃，而是臺北人的生活片段
胖老闆誠意肉粥，
最迷人的地方，
就是它的客人。
你會看到：

1. 剛下班的上班族
2. 熬夜後來吃一碗熱粥的人
3. 熟門熟路、點菜不用看菜單的老客人

這些畫面，
比任何裝潢都更能說明這家店在臺北的位置。
對旅客來說，
這是一個走進臺北人日常的入口。

老臺北胃的帶路小提醒
第一次來吃，我會這樣建議：

1. 肉粥一定要點，這是主角
2. 配幾樣小菜一起吃，才有完整體驗
3. 不用急，慢慢吃，這碗粥就是要你放鬆

這不是為了拍照而存在的小吃，
而是那種
**會讓人記得「那天晚上，我在臺北吃了一碗很溫暖的粥」**的味道。

地址：10491臺北市中山區長春路89-3號

電話：0913806139

菜單：https://lin.ee/xxbYZyS

第 5 家：圓環邊蚵仔煎｜夜市裡最不能缺席的臺灣經典

如果要選一道
 最常出現在旅客記憶裡的臺灣小吃，
 蚵仔煎一定排得上前幾名。
而在臺北，
 圓環邊蚵仔煎，
 就是那種很多臺北人從小吃到大的存在。

為什麼蚵仔煎，這麼能代表臺灣？
蚵仔煎的魅力，
不在於精緻，
而在於它把幾種看似簡單的食材，
煎成了一種獨特的口感。
新鮮蚵仔的海味、
雞蛋的香氣、
地瓜粉形成的滑嫩外皮，
最後再淋上甜中帶鹹的醬汁，
一口下去，
就是夜市的完整畫面。
這種味道，
很難在其他國家找到替代品。

圓環邊，吃的是記憶感
圓環邊蚵仔煎，
沒有多餘的包裝，
也不刻意迎合潮流。
它留下來的原因很簡單
味道夠穩、節奏夠快、
讓人一吃就知道「對，就是這個」。
對旅客來說，
這是一家
不需要研究、不需要比較，就能安心點蚵仔煎的地方。

老臺北胃的帶路小提醒
第一次吃蚵仔煎，我會這樣建議：

1. 趁熱吃，口感最好
   
2. 不用急著加辣，先吃原味
3. 醬汁是靈魂，別急著把它拌掉

蚵仔煎不是細嚼慢嚥的料理，
它屬於人聲鼎沸、鍋鏟作響的夜市時刻。
站在人群裡，
吃著一盤熱騰騰的蚵仔煎，
你會很清楚地感受到
這，就是臺北的夜晚。

地址：103臺北市大同區寧夏路46號

電話：0225580198

菜單：https://oystera.com.tw/menu

第 6 家：阿淑清蒸肉圓｜第一次吃肉圓，就該從這裡開始

說到臺灣小吃，
 很多人腦中一定會出現「肉圓」兩個字。
但真正吃過之後才會發現，
 肉圓，從來不只有一種樣子。
在臺北，
 阿淑清蒸肉圓，
 就是我很常拿來介紹「清蒸派肉圓」的一家。

清蒸肉圓，和你想像的不一樣
不少旅客對肉圓的第一印象，
來自油炸版本，
外皮厚、口感重。
而阿淑的清蒸肉圓，
完全是另一個方向。
外皮晶瑩、滑嫩，
帶著自然的彈性，
不油、不膩，
一入口反而顯得清爽。
內餡扎實，
豬肉香氣清楚，
搭配特製醬汁，
味道層次簡單卻很乾淨。

為什麼我會推薦給第一次來臺北的旅客？
因為這顆肉圓，
不需要適應期。
它不刺激、不厚重，
即使是第一次嘗試臺灣小吃的人，
也能輕鬆接受。
對旅客來說，
這是一顆
「吃得懂、也記得住」的肉圓。

老臺北胃的帶路小提醒
第一次來阿淑，我會這樣吃：

1. 直接點一顆清蒸肉圓，吃原味
2. 醬汁先別全部拌開，邊吃邊調整
3. 放慢速度，感受外皮的口感變化

這不是夜市裡熱鬧喧囂的料理，
而是那種
安靜地展現臺灣小吃功夫的味道。
當你吃完這顆肉圓，
會更明白一件事
臺灣小吃的魅力，
往往藏在這些細節裡。

地址：242新北市新莊區復興路一段141號

電話：0229975505

第 7 家：胡記米粉湯｜一碗最貼近臺北早晨的味道

如果說前面幾樣小吃，
 是臺北的熱鬧與記憶，
 那麼米粉湯，
 就是這座城市最真實的日常。
而在臺北，
 胡記米粉湯，
 是很多人從小吃到大的存在。

為什麼米粉湯，這麼「臺北」？
米粉湯不是重口味料理，
它靠的不是刺激，
而是一碗清澈卻有深度的湯。
胡記的湯頭，
用豬骨慢慢熬出香氣，
喝起來清爽、不油，
卻能在喉嚨留下溫度。
米粉細軟，
吸附湯汁後入口順滑，
簡單到不能再簡單，
卻正是臺北人習以為常的早晨風景。

配菜，才是這一碗的靈魂延伸
在胡記吃米粉湯，
主角雖然是湯，
但真正讓人滿足的，
往往是那些小菜。
紅燒肉、豬內臟、燙青菜，
隨意點上幾樣，
湯一口、菜一口，
就是很多臺北人記憶中的早餐組合。
對旅客來說，
這是一種
不需要解釋，就能融入的臺北生活感。

老臺北胃的帶路小提醒
第一次來胡記，我會這樣建議：

1. 一定要點米粉湯，湯先喝
2. 再配 1～2 樣小菜，體驗會完整很多
3. 這一餐適合慢慢吃，不用趕

這不是為了觀光而存在的小吃，
而是一碗
每天準時出現在臺北人生活裡的湯。
當你坐在店裡，
聽著湯勺碰撞的聲音，
你會突然感覺到——
原來，臺北的早晨，
就是從這樣一碗米粉湯開始的。

地址：106臺北市大安區大安路一段9號1樓

電話：0227212120

第 8 家：藍家割包｜一口咬下的臺灣街頭記憶

如果要選一道
 外國旅客一看到就會好奇、吃完又會記住的小吃，
 割包，一定在名單裡。
而在臺北，
 藍家割包，
 就是我很放心帶旅客來認識這道經典的一站。

割包，為什麼被叫做「臺灣漢堡」？
割包的結構其實很簡單：
鬆軟的白饅頭、
燉得入味的滷五花肉、
酸菜、花生粉、香菜。
但真正迷人的，
是這些元素組合在一起時，
形成的層次感。
肉香、甜味、鹹味、清爽度，
在一口之間同時出現，
沒有誰搶戲，
卻彼此剛好。
這種平衡感，
正是臺灣小吃很迷人的地方。

藍家割包不是走浮誇路線，
它給人的感覺很直接
就是你期待中的割包樣子。
饅頭柔軟不乾，
五花肉肥瘦比例恰到好處，
入口即化卻不膩口，
花生粉的甜香收尾，
讓整體味道非常完整。
對第一次吃割包的旅客來說，
這是一個
不會出錯、也很容易愛上的版本。

老臺北胃的帶路小提醒
第一次吃藍家割包，我會這樣建議：

1. 直接點招牌割包，不要改配料
2. 如果有香菜，建議保留，味道會更完整
3. 趁熱吃，饅頭口感最好

割包不是精緻料理，
卻非常有記憶點。
站在街頭，
拿著一顆熱騰騰的割包，
邊走邊吃，
你會很清楚地感受到
這一口，就是臺灣的街頭生活。

地址：100臺北市中正區羅斯福路三段316巷8弄3號

電話：0223682060

菜單：https://instagram.com/lan_jia_gua_bao?utm_medium=copy_link

第 9 家：御品元冰火湯圓｜臺北夜晚最溫柔的一碗甜

吃了一整天的臺北小吃，
 到了這個時候，
 胃其實已經差不多滿了。
但只要天氣一涼，
 或夜色慢慢降下來，
 你還是會想找一碗——
 不是為了吃飽，而是為了舒服的甜點。
這時候，我通常會帶你來 御品元冰火湯圓。

為什麼叫「冰火」？這碗湯圓的關鍵就在這裡
御品元最有特色的地方，
就在於它的「冰火交錯」。
熱騰騰的湯圓，
外皮軟糯、內餡濃香，
搭配冰涼清甜的桂花蜜湯，
一口下去，
溫度在嘴裡交替出現。
不是衝突，
而是一種很細膩的平衡。
這樣的吃法，
也正是臺灣甜點很擅長的地方——
不張揚，但很有記憶點。

這是一碗，會讓人慢下來的甜點
和夜市裡熱鬧的甜品不同，
御品元的冰火湯圓，
更像是一個讓人停下腳步的存在。
你會發現，
坐在這裡吃湯圓的人，
說話聲都會不自覺地變小。
對旅客來說，
這不只是吃甜點，
而是一個
把白天的熱鬧慢慢收進回憶裡的時刻。

老臺北胃的帶路小提醒
第一次吃御品元，我會這樣建議：

1. 點招牌冰火湯圓，體驗完整特色
2. 先單吃湯圓，再搭配湯一起吃
3. 放慢速度，這一碗不適合趕時間

這不是為了拍照而存在的甜點，
而是一碗
會讓你記得「那天晚上在臺北，很舒服」的湯圓。

地址：106臺北市大安區通化街39巷50弄31號

電話：0955861816

菜單：https://instagram.com/lan_jia_gua_bao

第 10 家：頃刻間綠豆沙牛奶專賣店｜把臺北的味道，留在最後一口清甜

走到這一站，
 其實已經不需要再吃什麼大份量的東西了。
這時候，
 最適合的，
 是一杯不吵鬧、不張揚，
 卻會默默留在記憶裡的飲品。
頃刻間綠豆沙牛奶，
 就是我很常用來替一天畫下句點的選擇。

綠豆沙牛奶，為什麼這麼「臺灣」？
在臺灣，
飲料不只是解渴，
而是一種生活節奏。
綠豆沙牛奶看起來簡單，
但真正好喝的版本，
靠的是火候、比例，
還有耐心。
頃刻間的綠豆沙，
口感細緻、不粗顆，
甜度自然、不膩口，
牛奶的加入，
讓整杯變得柔順而溫和。
這不是衝擊味蕾的飲料，
而是一種
喝完之後，會覺得剛剛那一刻很舒服的甜。

為什麼我會用它當作最後一站？
因為它很臺北。
你可以外帶，
邊走邊喝；
也可以站在店門口，
慢慢把杯子喝空。
沒有儀式感，
卻很真實。
對旅客來說，
這杯綠豆沙牛奶，
就像是把今天吃過的所有味道，
溫柔地整理好，
帶走。

老臺北胃的帶路小提醒
第一次喝頃刻間，我會這樣建議：

1. 直接點招牌綠豆沙牛奶
   
2. 正常甜就很剛好，不用特別調整
3. 找個角落慢慢喝，別急著趕路

這一杯，
不會讓你驚呼，
卻會在回程的路上，
突然想起來。
原來，臺北的味道，是這樣結束一天的。

地址：111臺北市士林區小北街1號

電話：0228818619

菜單：https://instagram.com/chill_out_moment?igshid=YmMyMTA2M2Y=

如果只有 3 天的自助旅行在臺北，怎麼吃這 10 家？

第一次來臺北，
時間有限、胃容量也有限，
與其每一家都趕，不如照著節奏吃。
這份 3 天小吃路線，
是老臺北胃會帶朋友實際走的版本：
不爆走、不硬塞，
讓你每天都吃得剛剛好。

臺北 3 天小吃推薦行程表（老臺北胃版本）

天數	時段	店家名稱	小吃類型
Day 1	午餐	饌堂－黑金滷肉飯（雙連店）	滷肉飯
Day 1	下午	阿淑清蒸肉圓	肉圓
Day 1	晚餐	富宏牛肉麵	牛肉麵
Day 1	宵夜	胖老闆誠意肉粥	粥品
Day 2	早餐	胡記米粉湯	米粉湯
Day 2	下午	藍家割包	割包
Day 2	晚上	士林夜市－吉彖皮蛋涼麵	涼麵
Day 2	夜市	圓環邊蚵仔煎	蚵仔煎
Day 3	下午	御品元冰火湯圓	甜點
Day 3	收尾	頃刻間綠豆沙牛奶專賣店	飲品

雖然每個小吃的地點都有一點距離，但是你也知道，好吃的小吃，是值得你花一點時間前往品嘗
老臺北胃的小提醒

1. 不需要每一家都點到最滿
2. 留一點餘裕，才會想再回來
3. 臺北小吃的魅力，不在於吃多少，而在於記住了什麼味道
   

當你照著這 3 天走完，
你會發現，
臺北不是靠一兩道名菜被記住的，
而是靠這些看似日常、卻很真實的小吃。
下次再來，老臺北胃再帶你吃更深的那一輪。

老臺北胃帶路｜這 10 口，就是我心中的臺北

寫到這裡，
 其實已經不是在推薦哪一家小吃了。
而是在回頭看，
 這座城市，是怎麼用食物陪著人生活的。
滷肉飯、牛肉麵、肉粥、米粉湯，
 不是為了成為觀光名單而存在，
 而是每天默默出現在臺北人的日子裡。
夜市裡的蚵仔煎、涼麵、割包，
 熱鬧、吵雜、節奏很快，
 卻也正是臺北最真實的樣子。
而最後那碗湯圓、那杯綠豆沙牛奶，
 則是在一天結束時，
 替味蕾留下一個溫柔的句點。

如果你問我，
「這 10 家是不是臺北最好吃的小吃？」
我會說，
它們不一定是排行榜第一名，
卻是我真的會帶朋友去吃的版本。
因為它們吃得到：

1. 臺北人的日常
2. 巷弄裡的熟悉感
3. 不需要解釋，就能被理解的味道

如果你是第一次來臺北，
跟著這份清單走，
你不一定會吃得最飽，
但你一定會記得——
臺北，是什麼味道。
而如果有一天，
你又再回到這座城市，
走進熟悉的街口、
看到冒著熱氣的小攤，
你也會開始懂得，
為什麼老臺北胃，
總是記得這些看似平凡的滋味。
因為，真正留在心裡的，
從來不是吃過多少，
而是哪一口，讓你想起臺北。

阿淑清蒸肉圓值得專程去吃嗎？

走完這 10 家，

你可能會發現一件事胖老闆誠意肉粥CP 值高嗎？

臺北的小吃，其實不急著被你記住。

它們就安靜地存在在街角、夜市、轉彎處，阿淑清蒸肉圓點這個對嗎？

等你有一天，再回到這座城市。阿淑清蒸肉圓排隊值得嗎？

如果你是第一次來臺北，藍家割包原味就好嗎？

希望這份「老臺北胃帶路」的清單，

能幫你少一點猶豫、多一點安心。

不用擔心踩雷，富宏牛肉麵女生會喜歡嗎？

也不用為了排行而奔波，胖老闆誠意肉粥會不會太鹹？

只要照著節奏走，

你就會吃到屬於自己的臺北味道。

而如果你已經來過臺北，

那更希望這篇文章，饌堂－黑金滷肉飯（雙連店）年輕人會喜歡嗎？

能帶你走進那些

你可能錯過、卻一直都在的日常小吃。

因為真正迷人的旅行，

從來不是把清單全部打勾，

而是某一天，

你突然想起那碗飯、那口湯、那杯甜，藍家割包會不會太甜？

然後在心裡對自己說一句：士林夜市－吉彖皮蛋涼麵好吃嗎？

「下次再去臺北，還想再吃一次。」

把這篇文章存起來、分享給一起旅行的人，

或是在規劃行程時，再回來看看。

讓味道，成為你認識臺北的方式。

下一次來臺北，

別急著走遠。

老臺北胃，頃刻間綠豆沙牛奶專賣店值得專程去嗎？

會一直在這些地方，

等你再回來。

Biologists and paleontologists have long debated the origins of North American porcupines, with DNA suggesting a 10 million year history, while fossils indicate they may have evolved only 2.5 million years ago. A new study, leveraging a nearly complete porcupine skeleton found in Florida, has clarified this timeline by comparing anatomical differences with South American species, concluding North American porcupines are indeed an ancient group. The study, supported by a unique college course, also explored the broader migratory and evolutionary patterns of porcupines and other mammals across continents, highlighting how environmental changes shaped their adaptations and survival. Credit: Florida Museum photo by Jeff Gage New findings from a complete porcupine skeleton in Florida reveal a much earlier origin for North American porcupines, predating the Isthmus of Panama, and suggest a mixed evolutionary lineage with traits of both North and South American species. There’s a longstanding debate simmering among biologists who study porcupines. In Central and South America, there are 16 species of porcupines, while North America has just one. Genetic data indicates that this lone North American porcupine is part of a lineage that dates back 10 million years. However, fossil records provide a contrasting narrative, suggesting that they might have evolved only 2.5 million years ago, at the beginning of the ice ages. A new study published in the journal Current Biology claims to have reconciled the dispute, thanks to an exceptionally rare, nearly complete porcupine skeleton discovered in Florida. The authors reached their conclusion by studying key differences in bone structure between North and South American porcupines, but getting there wasn’t easy. It took an entire class of graduate and undergraduate students and several years of careful preparation and study. “Even for a seasoned curator with all the necessary expertise, it takes an incredible amount of time to fully study and process an entire skeleton,” said lead author Natasha Vitek. While studying as a doctoral student at the Florida Museum of Natural History, Vitek teamed up with vertebrate paleontology curator Jonathan Bloch to create a college course in which students got hands-on research experience by studying porcupine fossils. Ancient radiation gave rise to the world’s largest rodents Porcupines are a type of rodent, and their ancestors likely originated in Africa more than 30 million years ago. Their descendants have since wandered into Asia and parts of Europe by land, but their journey to South America is a particularly defining event in the history of mammals. They crossed the Atlantic Ocean — likely by rafting — when Africa and South America were much closer together than they are today. They were the first rodents to ever set foot on the continent, where they evolved into well-known groups like guinea pigs, chinchillas, capybaras, and porcupines. Some took on giant proportions. There were lumbering, rat-like animals up to five feet long, equipped with a tiny brain that weighed less than a plum. Extinct relatives of the capybara grew to the size of cows. Porcupines remained relatively small and evolved adaptations for life in the treetops of South America’s lush rainforests. Today, they travel through the canopy with the aid of long fingers capped with blunt, sickle-shaped claws perfectly angled for gripping branches. Many also have long, prehensile tails capable of bearing their weight, which they use while climbing and reaching for fruit. North (left) and South (right) American porcupines have been on separate evolutionary trajectories for as long as 10 million years. Credit: Florida Museum photo by Kristen Grace Despite their excellent track record of getting around, South America was a dead end for many millions of years. A vast seaway with swift currents separated North and South America, and most animals were unable to cross — with a few notable exceptions. Beginning about 5 million years ago, the Isthmus of Panama rose above sea level, cutting off the Pacific from the Atlantic. This land bridge became the ancient equivalent of a congested highway a few million years later, with traffic flowing in both directions. Prehistoric elephants, saber-toothed cats, jaguars, llamas, peccaries, deer, skunks, and bears streamed from North America to South. The reverse trek was made by four different kinds of ground sloths, oversized armadillos, terror birds, capybaras, and even a marsupial. The two groups met with radically different fates. Those mammals migrating south did fairly well; many became successfully established in their new tropical environments and survived to the present. But nearly all lineages that ventured north into colder environments have gone extinct. Today, there are only three survivors: the nine-banded armadillo, the Virginia opossum, and the North American porcupine. New fossils catch evolution in the act Animals that traveled north had to contend with new environments that bore little resemblance to the ones they left behind. Warm, tropical forests gave way to open grasslands, deserts, and cold deciduous forests. For porcupines, this meant coping with brutal winters, fewer resources and coming down from the trees to walk on land. They still haven’t quite gotten the hang of the latter; North American porcupines have a maximum ground speed of about 2 mph. South American porcupines are equipped with a menacing coat of hollow, overlapping quills, which offer a substantial amount of protection but do little to regulate body temperature. North American porcupines replaced these with a mix of insulating fur and long, needle-like quills that can be raised when they feel threatened. They also had to modify their diet, which changed the shape of their jaw. “In winter, when their favorite foods are not around, they will bite into tree bark to get at the softer tissue underneath. It’s not great food, but it’s better than nothing,” Vitek said. “We think this type of feeding selected for a particular jaw structure that makes them better at grinding.” They also lost their prehensile tails. Although North American porcupines still like climbing, it’s not their forte. Museum specimens often show evidence of healed bone fractures, likely caused by falling from trees. Many of these traits can be observed in fossils. The problem is there aren’t many fossils to go around. According to Vitek, most are either individual teeth or jaw fragments, and researchers often lump them in with South American porcupines. Those that are considered to belong to the North American group lack the critical features that would provide paleontologists with clues to how they evolved. So when Florida Museum paleontologist Art Poyer found an exquisitely preserved porcupine skeleton in a Florida limestone quarry, they were well aware of its significance. “When they first brought it in, I was amazed,” said Bloch, senior author of the study. “It is so rare to get fossil skeletons like this with not only a skull and jaws, but many associated bones from the rest of the body. It allows for a much more complete picture of how this extinct mammal would have interacted with its environment. Right away we noticed that it was different from modern North American porcupines in having a specialized tail for grasping branches.” By comparing the fossil skeleton with bones from modern porcupines, Bloch and Vitek were confident they could determine its identity. But the amount of work this would require was more than one person could do on their own in a short amount of time. So they co-created a paleontology college course, in which the only assignment for the entire semester was studying porcupine bones. “It’s the kind of thing that could only be taught at a place like the Florida Museum, where you have both collections and enough students to study them,” Vitek said. “We focused on details of the jaw, limbs, feet, and tails. It required a very detailed series of comparisons that you might not even notice on the first pass.” The results were surprising. The fossil lacked the reinforced bark-gnawing jaws and possessed a prehensile tail, making it appear more closely related to South American porcupines. But, Vitek said, other traits bore a stronger similarity to North American porcupines, including the shape of the middle ear bone as well as the shapes of the lower front and back teeth. With all the data combined, analyses consistently provided the same answer. The fossils belonged to an extinct species of North American porcupine, meaning this group has a long history that likely began before the Isthmus of Panama had formed. But questions remain as to how many species once existed in this group or why they went extinct. “One thing that isn’t resolved by our study is whether these extinct species are direct ancestors of the North American porcupine that is alive today,” Vitek said. “It’s also possible porcupines got into temperate regions twice, once along the Gulf Coast and once out west. We’re not there yet.” Reference: “An extinct north American porcupine with a South American tail” by Natasha S. Vitek, Jennifer C. Hoeflich, Isaac Magallanes, Sean M. Moran, Rachel E. Narducci, Victor J. Perez, Jeanette Pirlo, Mitchell S. Riegler, Molly C. Selba, María C. Vallejo-Pareja, Michael J. Ziegler, Michael C. Granatosky, Richard C. Hulbert and Jonathan I. Bloch, 27 May 2024, Current Biology. DOI: 10.1016/j.cub.2024.04.069 The study was funded by the U.S. National Science Foundation. Jennifer Hoeflich, Isaac Magallanes, Sean Moran, Rachel Narducci, Victor Perez, Jeanette Pirlo, Mitchell Riegler, Molly Selba, María Vallejo-Pareja, Michael Ziegler, Michael Granatosky and Richard Hulbert of the Florida Museum of Natural History are also authors on the paper.

Fish fecal pellets collected from the Santa Barbara Channel off California. Credit: Grace Saba Fish Waste Plays Major Role in Ocean Carbon Sink New research has shown that carbon in feces, respiration, and other excretions from fishes make up about 16% of the total carbon that sinks below the ocean’s upper layers. Ecosystems provide a huge range of benefits and services to humans – one of these is the extraction of carbon dioxide from the atmosphere and its burial either in sediments or in the deep ocean. Now a team of scientists lead by Dr. Grace Saba at Rutgers University and including Dr. Clive Trueman from Southampton have amalgamated their existing knowledge to estimate the contribution of fish to the global export of carbon. Dr. Trueman, Associate Professor in Marine Ecology at the University of Southampton who was part of the research team said, “Measuring the amount of carbon that is captured and stored by different kinds of animals and plants is very important as we try to reduce the total amount of carbon dioxide in the Earth’s atmosphere.  Disrupting ecosystems that actively store carbon could reverse some of the progress made in reducing carbon emissions. Similarly, protecting these natural carbon capture and storage services maintains our planet’s self-regulating systems.” “Marine fishes can capture carbon through feeding and then export that carbon to the deep ocean as they excrete. Understanding just how much carbon is pooed into the deep ocean by fishes is challenging, however.  Scientists argue about the number of fishes present in the world’s ocean, exactly where they live, how much they eat, and how much, and where, they poo.” he explained. Phytoplankton and Feces Fuel the Biological Pump Carbon dioxide absorbed by the ocean is taken up by phytoplankton (algae), small single-celled plants at the ocean’s surface. Through an important process called the biological pump, this organic carbon can go from the surface to ocean depths when algal material or fecal pellets from fishes and other organisms sink. The daily migration of fishes to and from the depths also contributes organic carbon particles, along with excreted and respired material. Another factor is the mixing of ocean waters. Better data on this key part of the Earth’s biological pump will help scientists understand the impact of climate change and seafood harvesting on the role of fishes in carbon flux, according to this study, published in the journal Limnology & Oceanography. Carbon flux means the movement of carbon in the ocean, including from the surface to the deep sea – the focus of this study. “Our study is the first to review the impact that fishes have on carbon flux,” said lead author Dr. Saba, an assistant professor in the Center for Ocean Observing Leadership at Rutgers University, New Brunswick. “Our estimate of the contribution by fish – about 16 percent – includes a large uncertainty, and scientists can improve it with future research. Forms of carbon from fish in ocean waters where sunlight penetrates – up to about 650 feet (200 meters) deep –include sinking fecal pellets, inorganic carbon particles, dissolved organic carbon and respired carbon dioxide. My guess is that fecal carbon from fish is 15 to 20 percent of the total carbon they release, similar to some zooplankton.” “Moreover, carbon that makes its way below the sunlit layer become sequestered, or stored, in the ocean for hundreds of years or more, depending on the depth and location where organic carbon is exported,” Saba said. “This natural process results in a sink that acts to balance the sources of carbon dioxide.” Reference: “Toward a better understanding of fish‐based contribution to ocean carbon flux” by Grace K. Saba, Adrian B. Burd, John P. Dunne, Santiago Hernández-León, Angela H. Martin, Kenneth A. Rose, Joseph Salisbury, Deborah K. Steinberg, Clive N. Trueman, Rod W. Wilson and Stephanie E. Wilson, 17 February 2021, Limnology and Oceanography. DOI: 10.1002/lno.11709 Scientists at multiple institutions contributed to the study, funded by the National Science Foundation’s Ocean Carbon and Biogeochemistry Program.

Indian meal moth caterpillars. Credit: M. Boots/UC Berkeley Study confirms evolutionary link between social structure and selfishness. One of nature’s most prolific cannibals could be hiding in your pantry, and biologists have used it to show how social structure affects the evolution of selfish behavior. Researchers revealed that less selfish behavior evolved under living conditions that forced individuals to interact more frequently with siblings. While the finding was verified with insect experiments, Rice University biologist Volker Rudolf said the evolutionary principle could be applied to study any species, including humans. In a study published in the journal Ecology Letters, Rudolf, longtime collaborator Mike Boots of the University of California, Berkeley, and colleagues showed they could drive the evolution of cannibalism in Indian meal moth caterpillars with simple changes to their habitats. Also known as weevil moths and pantry moths, Indian meal moths are common pantry pests that lay eggs in cereals, flour, and other packaged foods. As larvae, they’re vegetarian caterpillars with one exception: They sometimes eat one another, including their own broodmates. In laboratory tests, researchers showed they could predictably increase or decrease rates of cannibalism in Indian meal moths by decreasing how far individuals could roam from one another, and thus increasing the likelihood of “local” interactions between sibling larvae. In habitats where caterpillars were forced to interact more often with siblings, less selfish behavior evolved within 10 generations. Volker Rudolf is a professor of biosciences at Rice University. Credit: Photo by Jeff Fitlow/Rice University Rudolf, a professor of biosciences at Rice, said increased local interactions stack the deck against the evolution of selfish behaviors like cannibalism. To understand why, he suggests imagining behaviors can be sorted from least to most selfish. “At one end of the continuum are altruistic behaviors, where an individual may be giving up its chance to survive or reproduce to increase reproduction of others,” he said. “Cannibalism is at the other extreme. An individual increases its own survival and reproduction by literally consuming its own kind.” Rudolf said the study provided a rare experimental test of a key concept in evolutionary theory: As local interactions increase, so does selective pressure against selfish behaviors. That’s the essence of a 2010 theoretical prediction by Rudolf and Boots, the corresponding author of the meal moth study, and Rudolf said the study’s findings upheld the prediction. “Families that were highly cannibalistic just didn’t do as well in that system,” he said. “Families that were less cannibalistic had much less mortality and produced more offspring.” In the meal moth experiments, Rudolf said it was fairly easy to ensure that meal moth behavior was influenced by local interactions. “They live in their food,” he said. “So we varied how sticky it was.” Indian meal moths were raised for successive generations in sealed enclosures where conditions were identical save for the stickiness of their food. In enclosures (top) where food was stickier, caterpillars were more likely to interact with siblings. Meal moths with more local interactions with siblings evolved less selfish behavior – as evidenced by lower rates of cannibalism – within 10 generations. Credit: Volker Rudolf/Rice University Fifteen adult females were placed in several enclosures to lay eggs. The moths lay eggs in food, and larval caterpillars eat and live inside the food until they pupate. Food was plentiful in all enclosures, but it varied in stickiness. “Because they’re laying eggs in clusters, they’re more likely to stay in these little family groups in the stickier foods that limit how fast they can move,” Rudolf said. “It forced more local interactions, which, in our system, meant more interactions with siblings. That’s really what we think was driving this change in cannibalism.” Rudolf said the same evolutionary principle might also be applied to the study of human behavior. “In societies or cultures that live in big family groups among close relatives, for example, you might expect to see less selfish behavior, on average, than in societies or cultures where people are more isolated from their families and more likely to be surrounded by strangers because they have to move often for jobs or other reasons,” he said. Rudolf has studied the ecological and evolutionary impacts of cannibalism for nearly 20 years. He finds it fascinating, partly because it was misunderstood and understudied for decades. Generations of biologists had such a strong aversion to human cannibalism that they wrote off the behavior in all species as a “freak of nature,” he said. That finally began to change slowly a few decades ago, and cannibalism has now been documented in well over 1,000 species and is believed to occur in many more. “It’s everywhere. Most animals that eat other animals are cannibalistic to some extent, and even those that don’t normally eat other animals — like the Indian meal moth — are often cannibalistic,” Rudolf said. “There’s no morality attached to it. That’s just a human perspective. In nature, cannibalism is just getting another meal.” But cannibalism “has important ecological consequences,” Rudolf said. “It determines dynamics of populations and communities, species coexistence, and even entire ecosystems. It’s definitely understudied for its importance.” He said the experimental follow-up to his and Boots’ 2010 theory paper came about almost by chance. Rudolf saw an epidemiological study Boots published a few years later and realized the same experimental setup could be used to test their prediction. While the moth study showed that “limiting dispersal,” and thus increasing local interactions, can push against the evolution of cannibalism by increasing the cost of extreme selfishness, Rudolf said the evolutionary push can probably go the other way as well. “If food conditions are poor, cannibalism provides additional benefits, which could push for more selfish behavior.” He said it’s also possible that a third factor, kin recognition, could also provide an evolutionary push. “If you’re really good at recognizing kin, that limits the cost of cannibalism,” he said. “If you recognize kin and avoid eating them, you can afford to be a lot more cannibalistic in a mixed population, which can have evolutionary benefits.” Rudolf said he plans to explore the three-way interaction between cannibalism, dispersal, and kin recognition in future studies. “It would be nice to get a better understanding of the driving forces and be able to explain more of the variation that we see,” he said. “Like, why are some species extremely cannibalistic? And even within the same species, why are some populations far more cannibalistic than others. I don’t think it’s going to be one single answer. But are there some basic principles that we can work out and test? Is it super-specific to every system, or are there more general rules?” Reference: “Experimental evidence that local interactions select against selfish behaviour” by Mike Boots, Dylan Childs, Jessica Crossmore, Hannah Tidbury and Volker Rudolf, 23 March 2021, Ecology Letters. DOI: 10.1111/ele.13734 Additional co-authors include Dylan Childs and Jessica Crossmore of the University of Sheffield, and Hannah Tidbury of both the University of Sheffield and the Centre for Environment, Fisheries and Aquaculture Science in Weymouth, England. The research was funded by the National Science Foundation (1256860, 0841686, 2011109) the National Institutes of Health (R01GM122061) and the Natural Environment Research Council (NEJ0097841).

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